Yali Guo1†
Yifei Luo2,3†
Yuting Ma2
Afito Luciano2Jingjing Huang1,4Ye Li1,4Shiwen Wang1,4Yuequn Niu1Zhou Li1
Jifeng Cai1,4,5*
Fanming Meng1,2,4*- 1Department of Forensic Medicine, School of Basic Medical Sciences, Xinjiang Medical University, Urumqi, Xinjiang, China
- 2Department of Parasitology, Central South University, Changsha, China
- 3Shaoyang Center for Disease Control and Prevention, Shaoyang, Hunan, China
- 4Key Laboratory of Forensic Medicine, Xinjiang Medical University, Urumqi, China
- 5Department of Forensic Science, School of Basic Medical Sciences, Central South University, Changsha, Hunan, China
Forensic entomology focuses on insects associated with decomposing remains to provide clues for forensic investigations. Among those insects, scuttle flies are uniquely capable of colonizing enclosed spaces and underground burial sites where other insects cannot access, often serving as the sole entomological “witnesses” to provide crucial evidence in forensic practice. This review highlights the forensic significance of scuttle flies, including the biological characters, diversity and behaviors of scuttle flies from forensic aspects based on reported cases. We investigate the biological and environmental factors influencing their utility in forensic investigation, and scuttle fly species commonly found on corpses were also summarized. Additionally, suggestions on future research directions of scuttle flies and how their biological characteristics can enhance their application in forensic entomology are also provided.
1 Introduction
Forensic entomology is a branch of applied entomology that uses the presence of necrophagous insects to assist in criminal investigations, such as by estimating the minimum post-mortem interval (PMImin), inferring changes in the corpse’s position, and so forth, to provide auxiliary evidence (1). In forensic research, the accurate assessment of the time of death, the postmortem interval (2), occupies an extremely critical position (3). Flies act as important forensic indicators as they are the foremost visitors of the corpse and act as silent witnesses after the accomplishment of a crime (2).
Forensically relevant insects (such as scuttle flies, flesh flies, and house flies) can rapidly locate and colonize a freshly deceased body. However, when the corpse is situated in a relatively enclosed environment (such as sealed containers, locked rooms, buried coffins) (4–6), a physical barrier is formed by theses conditions, which hinders their colonization. Based on this, two key forensic questions arise. First, the delay in egg-laying time (from several hours to several days or weeks) may lead to an underestimation of the PMI. Because traditional indicators based on insect development time do not take these barriers into account (7). Although isolated reports describe Phoridae insect associated with long-buried or skeletonized remains (8), such occurrences represent exceptional and highly specific conditions rather than typical Diptera colonization patterns. Diptera generally cannot oviposit or complete development on fully skeletonized bodies, where beetles dominate the fauna (9, 10). Second, these enclosed environments stabilize the decomposition process by reducing temperature fluctuations, maintaining constant humidity, reducing air flow, and weakening competition among insect species (11), which further modifies colonization dynamics and development rate of insects.
Scuttle flies are among the most species-rich and worldwide distributed groups of insects (12). Adult scuttle flies are small, ranging from 0.75~8.0 mm in size, and the larvae are 1~4 mm. Adults exhibit colors such as dark black, gray-brown, or yellowish. Adults have an enlarged thorax with a characteristic humpbacked appearance. Adults and larvae prefer moist environments and live on decaying carrion, carcass, dung, plants, flowers, fungi, beehives, or anthills (13).
Due to the smaller body size and the natural ability to locate food sources in a relatively confined environment, scuttle flies tend to be found in crime scenes that other insects cannot reach, which gives them special meaning during forensic investigation (14). Lutz et al. (15) found that 949 insect-related cases, 80% entomological evidence, can provide day-specific PMImin estimates. Table 1 summarizes the scuttle flies found on human corpses in forensic cases and their estimation of PMImin.
Table 1. Scuttle flies found on human corpses in forensic cases and their estimation of PMI.
2 The forensic application scenarios of Scuttle flies
2.1 Buried corpses
Phoridae species are among the most typically encountered taxa on decaying carcasses in the soil at various depths (30). Many cases have been reported of Phoridae being found on bodies in graves, and most of the bodies were found skeletonized (5, 24, 26). In a case, the phorid fly is the only insect found on the corpse of a murder victim buried (<1 m depth) in a heavy clay soil in the East of England (26). It was reported to colonize carcasses at a depth of 60 centimeters in less than two weeks post-burial (31). Megaselia scalaris Loew could enter through the smallest holes to reach the buried carcasses, due to its tiny body size (32). M. scalaris can also detect carrion buried in coffins at a depth of around 1.83metres and oviposit on it, providing a food source for the larvae that hatch from the eggs, supporting their development until pupation (33).
The rate of decomposition of buried bodies is highly dependent on the depth of the burial and the ambient temperature (34). In an extreme case, Conicera tibialis Schmitz can invade corpses buried for years and successfully complete at least one generation of reproduction, where a saponified corpse was buried in a zinc coffin at a depth of about 2 meters (8). Although exceptional cases exist, Phoridae generally cannot breed on fully skeletonized remains. Saponification of corpses can slow down the degree of corruption (35). Since it can provide sufficient physical resources for Phoridae and support multi-generation reproduction, their presence in buried bodies is considered part of the cadaveric fauna and is not a reliable indicator for postmortem interval estimation. In addition, insect invasion of the buried remains will also impact the decomposition rate and pattern (36). It should be considered when estimating the postmortem interval.
2.2 Wrapping corpses and closed space
Firstly, to make the evidence disappear, some suspects would place the corpse in a relatively closed environment such as a quilt, a vehicle, or a suitcase (17, 37, 38). Due to the influence of the containing environment, wrapping will prolong the process by which insects search for corpses and reduce the number of eggs laid. In this regard, different wrapping methods will cause differences in the time when flies first come into contact with the corpse. For instance, plastic bags, simulated luggage, and plastic bags plus suitcases will gradually prolong the time for flies to come into contact with corpses for the first time (39). This is mainly attributed to differences in material, thickness, and breathability. While wrapping caused a 1 to 13 days delay in the arrival of other species, Phoridae did not exhibit any delay in reaching the carcass (40). In addition, differences in the exposure level of the corpse, its location, as well as other aspects of the discovery scene, will further influence the colonization process of the flies by altering the ease with which they come into contact with the corpse (39, 41).This phenomenon is consistent with the research of Andrews et al. (7), Phoridae were most abundant in the luggage, as opposed to the wheeled bins and the exposed environment. Cases that further support this view include: Ivorra et al. (29) found Puliciphora. rufipes Silva Figueroa on a body wrapped in a blanket. Similarly, Zuha et al. (42) reported that Puliciphora Dahl (Diptera: Phoridae) was found on rabbit carcasses placed in luggage and garbage. Besides, they also recorded that Dahliphora sigmoides Schmitz, Gymnoptera simplex Brues, M. scalaris, Puliciphora borinquenensis Wheeler, Puliciphora obtecta Meijere, and Spiniphora sp. were found on rabbit carcasses confined to sealed plastic bins (43). Due to the dual interference of insect colonization time and decomposition rate, this eventually leads to an impact on the accuracy of PMImin inference for wrapped corpses.
In addition to insect colonization, the wrapping also significantly alters the process of corpse decomposition. The decomposition time of package processing is longer than that of unpackage processing (44). This is due to plastic bags isolating the samples from the external environment, creating anaerobic conditions that promote fat decomposition and the formation of adipocere (35). Simultaneously, the impermeability of plastics leads to the accumulation of decomposition products during the degradation process, which slows down the decomposition rate (45). When comparing the corpses wrapped in plastic with those in an open environment, it was found that the duration of the fresh stage was the same for both, but the duration of the wrapped corpses in the remaining stages was longer (46). Because plastic wrapping creates a high-humidity and low-oxygen environment, which inhibits the activity of putrefactive bacteria and slows down tissue decomposition (47, 48). But it creates a high temperature environment and increases insect activity (49).
However, the influence of the characteristics of the wrapping material itself on decomposition is more complex. When comparing cotton fabric with plastic, cotton fabric causes local dryness due to its hygroscopic properties, delaying decomposition. It is worth noting that mulch made of natural fibers has been proven to accelerate the formation of liposis compared with that made of synthetic fibers in aquatic environments. The reason for this phenomenon is that natural fibers have a stronger water absorption capacity than synthetic fibers (50). This difference may stem from absorption. It may remove the decomposition products of the surface tissues of the corpse. This effect may disrupt the decomposition process and make the formation of fat more likely to occur.
Lastly, the interaction between the package and other environmental factors also needs to be considered. During the interaction between soil burial and wrapping, it was observed that the proportion of mass loss (the ratio of post-burial to pre-burial quality) indicated wrapping had a greater impact on the decomposition of corpses, but no impact on TBS (51).
2.3 Indoor building
Scuttle flies (Diptera: Phoridae), particularly M.scalaris, hold unique forensic significance in indoor death scene investigations. Unlike dominant necrophagous species like Lucilia sericata Meigen and Calliphora vicina Robineau-Desvoidy (Calliphoridae), Phoridae exhibit remarkable adaptability to enclosed environments (19, 52). Their tiny body size enables them to colonize cadavers in sealed indoor spaces through narrow gaps in doors or windows, even when these openings are only partially closed (18). This ecological trait explains their frequent dominance in indoor insect communities despite their relatively low abundance in regional surveys. For instance, while Phoridae accounted for merely 0.39% of the insect fauna in human cadavers across Tehran District (53), they were successfully documented in an indoor case within the same region, highlighting their niche specialization.
The vertical distribution patterns of scuttle flies further enhance their forensic utility. Although primarily associated with ground-level indoor cases (20), M. scalaris demonstrates exceptional resource exploitation capabilities in high-rise environments (6). Field experiments reveal they can colonize substrates at heights exceeding 100m through passive transport mechanisms and human-mediated dispersal (54), outperforming Calliphoridae and Sarcophagidae in vertical colonization efficiency (55). This behavioral adaptation positions Phoridae as critical PMImin indicators in high-rise death scenes where conventional blowfly evidence is absent. While current case reports linking Phoridae to elevated crime scenes remain limited (20), their predictable colonization patterns in confined spaces warrant increased attention from forensic entomologists. However, Phoridae are not strictly dependent on human cadavers; they can also develop in other substrates such as garbage, sewage pipes, and decaying plant materials. Therefore, samples collected from substrates associated with cadavers are applicable for PMI estimation.
2.4 Myiasis
Myiasis is an infestation of living humans or animals with dipterous larvae that feed on the host’s live or dead tissue, liquid matter, or ingested food from their host, and cause a wide range of infections depending on body location and the larvae’s relationship to the host (56). Necrophilic insect species found on human and animal corpses are also present in patients with myiasis. (57). Once the host dies and the tissues are fully decomposed, the same fly species may behave as necrophagous colonizers rather than as myiasis agents. Therefore, in forensic investigations it is important to distinguish true antemortem myiasis from postmortem colonization of cadavers by necrophagous larvae. M. rufipes, M. scalaris, and Megaselia spiracularis Schmitz were seen in cases of myiasis (58–60). Myiasis tends to show up in neglected cases. Under these conditions, the possibility of antenatal myiasis infection needs to be carefully considered when evaluating cases and PMImin (61). Diptera larvae can infest open wounds, predisposing the patient to severe secondary bacterial infection and, if untreated, even sepsis and death, which may indicate serious neglect or abuse (62). Critically, myiasis associated flies like M. scalaris exhibit ecological plasticity: they infest both living hosts and cadavers, necessitating caution when linking larvae solely to PMImin calculations. Standardized protocols should integrate wound microbiology, larval species identification, and developmental data to avoid misinterpreting antemortem infestations as postmortem colonization events.
3 Factors affecting scuttle flies on corpses
3.1 Temperature
The growth rate of insects varies with temperature. Studying the development rate of insects at different temperatures is conducive to predicting more accurate PMImin (63). Wang et al. (64) found that M. scalaris development duration decreased with increasing temperature in the range of 16-34°C. Thomas et al. (65) explored the growth rate of M. scalaris at 24°C, 28°Cand 32°C, respectively, increased by 32.1%, 13.9% and 45.5%, which indicates suitable temperatures promoting quicker development. Zuha and Omar (66) asuggested that M. scalaris larval length is possibly not ideal indicator for estimating larval age at temperatures less than 28°C and greater than 33°C. Feng et al. (67) found that the mean time of the pupation stage was negatively correlated with temperature for Dohrniphora cornuta Bigot. At lower temperatures and wetter conditions, adult lifespan increased for Phoridae (68).
Some species of Phoridae exhibit low temperature tolerance, and their adults can be active on snow surfaces at temperatures as low as -4°C (69, 70). Phoridae are active in the cold season, and most blowflies are inactive due to the low temperatures (25). Seasons can affect the distribution of phoridae indoors. Almutawa et al. (6) reported that adult phorids are only found indoors in winter. However, it was not found either indoors or outdoors in summer. While Phoridae maintain activity in winter indoor environments (18.0-19.6°C), elevated summer temperatures (38.1-43.0°C) may restrict their survival and colonization potential. Low temperature can cause developmental arrest. Megaselia giraudii Egger undergoes diapause during the pupal stage, with the emergence time extended by 184–187 days compared to normal conditions (71). Although current research has not directly confirmed the complete developmental history of Phoridae in winter or low-temperature environments. Therefore, the PMImin estimation, based on a comprehensive consideration of this biological characteristic, may have significant reference value when determining death cases in the cold season.
3.2 Humidity
Phoridae exhibit the highest abundance during the early decomposition stage (72). As an indicator species for this stage, this characteristic can serve as an auxiliary reference for inferring PMImin. During the fresh and bloated stages, the moisture content of the natural cadaveric openings might be as high as 70% to 80% (73), creating an ideal microenvironment for their activity. Under relatively humid conditions (> 75% RH), the lifespan of Phoridae will be prolonged (68). Experimental studies on M. scalaris demonstrate that substrate moisture significantly impacts pupal recovery and adult emergence. At 50% humidity, larvae complete development rapidly, whereas at 90% humidity, survival remains unchanged, but developmental time extends by approximately two days, and adult emergence rates drop by 20.64% (74). This is the reason why it is difficult for them to colonize over a long period of time. Furthermore, human cadaver moisture fluctuates across decomposition stages, directly affecting Phoridae colonization patterns. For forensic applications, this underscores the necessity of incorporating cadaver water content into PMImin estimates. Thus, when estimating PMImin for bodies with varying moisture levels, careful calibration based on humidity-driven developmental thresholds is essential to ensure reliability.
3.3 Burial depth and soil
Post-burial interval estimation, similar to PMImin, focuses on the development of data from insects detected on buried carcasses. The timing and pattern of insect colonization on buried corpses are determined by soil type and depth (75). Scuttle flies may arrive at corpses buried in deep soil in earlier phases of decomposition and may develop huge populations (76). Pastula and Merritt (31) reported that M. scalaris colonized corpses at a depth of 60 cm in the soil and was collected 7 days after burial, which can help estimate Post-burial interval (PBI) for forensic investigation. Moreover, it was discovered in excavated bodies buried in wooden coffins at a depth of 30–60 cm in the soil. AL-Mekhlafi et al. (77) found larvae of M. scalaris under 20 cm at the decay and advanced decay stage of decomposition on buried rabbits. Triphleba nudipalpis Becker appeared on the body of a murder victim buried in a meter of clay (26). Conicera similis Haliday was collected from a rat carcass buried 40 cm depth (78). C. tibialis can be found on corpses that have been buried at a depth of 2m for 18 years (8). Besides, soil moisture affects the growth and development of M. scalaris, the larval stage lasts the longest, and the pupation rate is the lowest in the soil with 0% water content (79). Therefore, when dealing with burial cases, it is also necessary to consider the impact of soil moisture on the growth and development of insects.
3.4 Drugs intake, pollutants, and toxic substances
Commonly, overdose-related deaths happen in isolated regions, particularly among lonely victims, frequently discovered in the latter stages of decomposition (80). Insects can be used as substitute evidence in poisoning death investigations (81). By analyzing larvae that feed on corpses, it is possible to identify the drugs present in decaying corpses (82). Insects colonize charred corpses, and poisons taken before death can be detected in the larvae (83).
Miller et al. (84) reported that M. scalaris pupae were discovered in the mummified body of a female drug addict who died 33 months earlier in her home. Several added drugs were found in the body. The pupae were found to contain amitriptyline and nortriptyline in the following analysis. The drug inhibits larval growth in a concentration-dependent manner (85). Clonazepam could expedite the growth of larvae, reaching their pupal stage earlier (86). M. scalaris that gnaw on carcasses in the presence of malathion experience reduced egg and pupa viability, diminished larval growth, and an extended larval stage, which can lead to errors in PMImin inference (87). In addition, a particular drug or poison can have different effects on different species (88). The drug’s effect on larval development might alter the agreement between the entomological PMImin estimation and the circumstantial facts in this situation. Current research indicates that there is no linear relationship between morphine concentrations in necrophagous insects and their larval feeding periods, as morphine does not undergo bioaccumulation within the insects (89). The efficacy of toxicological detection depends on whether the absorption rate of the drug in the host organisms exceeds its metabolic clearance rate. This dynamic equilibrium limits existing technologies to qualitative analyses of toxins in insect samples (e.g., confirming the presence or absence of specific substances) rather than enabling quantitative correlations with the postmortem interval (PMImin). Therefore, while toxicant detection holds significant referential value in forensic science, its sole application remains insufficient to support accurate estimation of the PMImin.
Because of the faster growth rate, the larval age may be overestimated. Alcaine-Colet et al. (90) reported that M. scalaris could complete its life cycle when fed on modeling clay, a substrate containing toxic inorganic compounds, which results in increased larval development time, reduced pupal development time, and a shorter adult life duration of M. scalaris. The effects of pollutants on the development of M. scalaris, such as hexavalent chromium and four species of selenium (91–93). Exploring insect toxicology can therefore estimate the minimum postmortem interval (94) and the circumstances of death in which toxic substances are suspected to have caused the death (88).
Interestingly, there are some reports of gasoline homicide. Gasoline can delay the time when scuttle flies enter the body. Rumiza et al. (95) used gasoline to mimic homicide or suicide. It has been established that the existence of gasoline and its odor on the corpse delays the decomposition process by postponing the entry of insects by 6 hours.
3.5 Nocturnal oviposition
Estimation of time has passed since death corresponds to the PMImin, which is based on the age of the oldest insects detected on bodies. Commonly, necrophagous flies like blowflies or flesh flies are inactive at night; hence, oviposition happens during the day (96). However, M. scalaris can lay eggs in complete darkness both during the day and at night, and the number of eggs laid at night is very similar to that laid under continuous dark conditions (97). If the insects in the discussion were nocturnal and laid eggs at night, it might cause the PMImin estimation to be off by up to 12 hours. Unlike Calliphoridae, common and representative research subjects in forensic entomology, which typically oviposit under sufficient daylight with extremely low egg production at night (98). M. scalaris can access the oviposition location and lay eggs in complete darkness both during the day and night. Nevertheless, the quantity of eggs laid during the day is larger. Moreover, the number of eggs laid during the night under LD conditions (12 hours of light, followed by 12 hours of darkness) is remarkably close to the amount in complete darkness (97). This implies that in situations where there are frequent periods of darkness, the number of eggs deposited is enough to support the population. It is advised that nocturnal oviposition should be included when using scuttle fly as a reference for PMImin estimation. Further, efforts to correlate numerous elements that contribute to nocturnal oviposition, such as temperature, relative humidity, and bait type, should be given.
3.6 Other factors
Dimorphism is an essential factor distinguishing between male and female development stages. The size dimorphism of M. scalaris is obvious during post-feeding and pupa stages, such as female post-feeding larvae and puparia are larger than males, and females have a longer total developmental period than males at almost all temperature ranges (66). It could help investigators estimate the development time of the species more accurately, thereby correctly estimating the development time of the species. Thus, the dimorphic difference should be considered when interpreting developmental data for PMImin analysis.
4 Scuttle flies of forensic interest
Phoridae is among the most biologically diverse and species-rich groups of insects, which contains 250 genera and about 3,400 species worldwide (12, 99). Table 1 summarizes the scuttle flies found on human corpses in forensic cases in terms of Phoridae species, distribution area, cadaver type, scene, and PMImin inference time.
4.1 Megaselia
M. scalaris is the most studied forensic-related scuttle fly for estimating the post-mortem interval of human remains (100). It is widely distributed and can be found in various habitats, including urban structures and tropical rainforest (101). It is a polyphagous species and can live on various foods, including decomposing organic materials, artificial media, and decaying substrate (102, 103). Additionally, M. scalaris is commonly associated with indoor death or neglect cases of humans or household animals (104).
This species played a significant role in decaying bodies located indoors or in concealed environments due to its small size and ability to penetrate these regions earlier than other insects (18, 21, 105). It is the only insect discovered breeding within a tightly closed seventh-floor flat in Japan (106). It was found on exposed pig carrion after 5–12 days of exposure (107). It is a common and typical forensic indicator that was found in the decay, advanced decay, and dry stages (32). Zhang et al. (108) found that the mean developmental duration of M. scalaris from egg to adult stage needs 417.7 hours at 25°C and an intrinsic optimum temperature of 21°C.
In a Phoridae study in Egypt, Megaselia curtineura Brues was the most abundant species, except M. scalaris (109). It is also recorded among the 16 species associated with 50 human remains in the tropical climate of northern Malaysia (110). Thevan et al. (21) first reported the use of the growth and development of M. curtineura for PMImin inference in forensic cases.
M. spiracularis had the longest development time (1131.1 h) at 16°C and the shortest development time (232.6 h) at 34°C (64). Although M. spiracularis and M. scalaris are closely related, their total developmental history at room temperature differs by 50.7 hours (64).
4.2 Dohrniphora
D. cornuta is a common species of Phoridae fly found indoors in China, and has also been reported in England, Spain, and Sweden, indicating that it is a widespread species (111, 112). Adults collected in a house in the Pennines (U.K.) at 380 meters above sea level were most likely attracted by a neighboring septic tank (113). The species was also collected on skeletonized human remains inside a closed tank (27). In addition, it developed faster at 24°C and 30°C, but was restricted beyond 36°C (67).
4.3 Conicera
C. tibialis, the coffin fly, is distinctive among the species commonly observed on buried bodies due to its peculiar association with cadavers in coffins and limited spaces. As a typical species adapted to the burial environment, C. tibialis can penetrate soil up to 2 meters deep to lay eggs and form multiple generations of reproduction in closed coffins (114). Martín-Vega et al. (8) further discovered that it could complete the life cycle on a corpse buried for 18 years. This species was discovered in a concrete coffin buried 1.8 m deep, 28 years after death (28). This breakthrough record not only reveals the ecological resilience of this species under extreme time spans but also challenges the boundaries of the traditional PMImin estimation model in forensic entomology.
C. similis has been found on both skeletonized human remains and decayed rat corpses, and these findings collectively confirm its ability to colonize dead bodies (27, 78).
4.4 Triphleba
T. nudipalpis is one of the most representative species. This species is found in the British Isles, continental Europe from Sweden to France, and Russia’s far east from the Czech Republic to Hungary (26). Larvae of T. nudipalpis feed on buried carcasses and are well-known soil breeders, as indicated by the large amount of phenology data acquired with emergence traps placed over the soil (115). Adults of T. nudipalpis have been reported visiting carrion (26). The larvae have been subsequently shown to be feeders on buried carcasses (116). Furthermore, they have been proven to be efficient in locating buried carrion, as is proven by the fact that pieces of flesh were buried at varying depths, and flies were reported as early as four days after burial (117). Conversely, many female phorids visit carrion for a protein-rich feed, which assists in the growth of their eggs, rather than laying mature eggs (118).
4.5 Puliciphora
Species of this genus are distributed worldwide and found primarily in warm-climate zones (119, 120). P. borinquenensis is a species of forensic importance in buried or closed environments (121). P. borinquenensis, P. obtecta, and Puliciphora beckeri Meijere were discovered on rabbit carcasses, as well as in luggage and trash cans (42). Puliciphora rufipes Silva Figueroa was found wrapped in a blanket (29).
5 Conclusion and perspectives
Compared to other insect species of forensic importance, scuttle flies have specific significance in forensic science, especially in cases involving bodies in enclosed spaces or environments. The tiny body size and necrophagous feeding habit may partly explain their advantage under such conditions. Traditional carbiophilic flies (such as those from the Calliphoridae family) can locate corpses because their antennae contain chemoreceptors and they are relatively sensitive to cadaverine and putrescine. Although the tiny size of the Phoridae is regarded as its main advantage in penetrating closed environments, the special sensitivity of its chemoreceptors may be underestimated. This mechanism has not yet been systematically verified in forensic practice. However, the biological mechanisms of why and how scuttle flies locate and invade corpses in enclosed spaces are not yet fully understood. In the future, it needs to be further explored in combination with molecular biology techniques (such as olfactory receptor gene analysis).
In enclosed spaces, the developmental patterns and oviposition behavior of scuttle flies are crucial for estimating PMImin (122). Previous research has collected data on the life stages of Phoridae, but much of this information originates from controlled laboratory settings with stable temperatures. However, this does not adequately reflect the complex environmental conditions encountered in real-world scenarios. Compared to open environments, factors such as temperature, humidity, and food availability within confined spaces are more stable, which affects the body decomposition and insect development in a unique manner. Moreover, the oviposition behavior of scuttle flies is also influenced by environmental conditions. Therefore, further exploration of the developmental patterns and oviposition behavior of Phoridae in simulated enclosed spaces is essential for improving the accuracy of PMImin estimation. In certain criminal scenarios, such as deep burial and chemical treatment of corpses, they may be the only reliable evidence. Looking to the future, it is necessary to promote the transition of Phoridae from “alternative tools” to “core tools”. This requires in-depth cooperation between the forensic medical community, entomologists, and data scientists to build a globally shared multi-omics database of Phoridae.
In recent years, advanced technological methods have been applied in forensic entomology field. To fully realize the potential of Phoridae as forensic indicators, future research must adopt advanced technological approaches currently available in forensic entomology. Key future directions include the application of multi-omics techniques to decipher their unique biological characteristics in concealed environments. Currently, machine learning algorithms have been used to model the developmental data of other necrophagous insects and cadaver microbiome succession independently for PMImin estimation (123–125). However, the integrated analysis of these two datasets remains a missing link. Constructing models that represent the entire “cadaver-Phoridae-microbiome” system represents a crucial objective for achieving the next key breakthrough in accurate PMImin estimation.
Scuttle flies are crucial in forensic entomology, especially in enclosed environments where other insects cannot access. Their small size and adaptability allow colonization of buried, wrapped, or indoor corpses, serving as unique “witnesses” for PMImin estimation. Factors like temperature, humidity, burial depth, and toxins influence their development and colonization, with species like M.scalaris demonstrating nocturnal oviposition and drug tolerance. Forensic cases worldwide highlight their role in extreme scenarios. Future research should explore molecular mechanisms of their environmental adaptation and build global multi-omics databases to enhance PMImin accuracy in concealed death scenes.
Author contributions
YG: Data curation, Investigation, Visualization, Writing – original draft. YFL: Data curation, Investigation, Visualization, Writing – original draft. YM: Data curation, Investigation, Writing – original draft. AL: Data curation, Investigation, Writing – original draft. JH: Data curation, Investigation, Writing – original draft. YL: Data curation, Investigation, Writing – original draft. SW: Data curation, Investigation, Writing – original draft. YN: Data curation, Investigation, Writing – original draft. ZL: Data curation, Investigation, Writing – original draft. JC: Conceptualization, Funding acquisition, Methodology, Supervision, Writing – review & editing. FM: Conceptualization, Funding acquisition, Methodology, Supervision, Writing – review & editing.
Funding
The author(s) declared that financial support was received for this work and/or its publication. This work was supported by the National Natural Science Foundation of China (grant number 32460248, 82471915, 32370554), the Natural Science Foundation of: Xinjiang Uygur Autonomous Region (No.2024D14016), and the Research and Innovation Team Project of Xinjiang Medical University (XYD2024C05).
Conflict of interest
The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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References
1. Wiraagni IA, Rezadhini M, Setiawan J, Sofyantoro F, Priyono DS, Septriani NI, et al. Research trends on forensic entomology for five decades worldwide. Leg Med (Tokyo). (2024) 71:102539. doi: 10.1016/j.legalmed.2024.102539
2. Babu BS, Kumar R, El-Sharnouby M, Salah M, Sayed S, Alhazmi A, et al. Calliphorids as forensic indicator to facilitate PMI estimation: A case study from Chhattisgarh, India. J King Saud Univ Sci. (2022) 34. doi: 10.1016/j.jksus.2021.101709
3. Bray S, Conlan X, and Harvey M. A preliminary evaluation of the utility of insects and algae for PMI estimation in confined, still-water environments. Aust J Forensic Sci. (2023) 55:129–40. doi: 10.1080/00450618.2021.1973100
4. Introna F, De Donno A, Santoro V, Corrado S, Romano V, Porcelli F, et al. The bodies of two missing children in an enclosed underground environment. Forensic Sci Int. (2011) 207:e40–7. doi: 10.1016/j.forsciint.2010.12.007
5. Mariani R, García-Mancuso R, Varela GL, and Kierbel I. New records of forensic entomofauna in legally buried and exhumed human infants remains in Buenos Aires, Argentina. J Forensic Legal Med. (2017) 52:215–20. doi: 10.1016/j.jflm.2017.09.012
6. Almutawa MY, Al-Khalifa MS, Al-Dhafer HM, Abdel-Dayem MS, Ebaid H, and Ahmed AM. Forensic investigation of carcass decomposition and dipteran fly composition over the summer and winter: a comparative analysis of indoor versus outdoor at a multi-story building. J Med Entomol. (2024) 61:877–90. doi: 10.1093/jme/tjae039
7. Andrews AG, Magni PA, and Dadour IR. The decomposition process in two restricted access environments in a temperate climate: Hard-covered suitcases and wheeled bins. Forensic Sci Int. (2025) 367:112368. doi: 10.1016/j.forsciint.2025.112368
8. Martín-Vega D, Gómez-Gómez A, and Baz A. The “coffin fly”Conicera tibialis (Diptera: Phoridae) breeding on buried human remains after a postmortem interval of 18 years. J forensic Sci. (2011) 56:1654–6. doi: 10.1111/j.1556-4029.2011.01839.x
9. Bala D and Singh N. Beetles and forensic entomology: A comprehensive review. J Entomological Res. (2015) 39:293. doi: 10.5958/0974-4576.2015.00036.5
10. Shao S, Liu S, Li L, Hu G, Zhang Y, and Wang Y. Research status of sarcosaprophagous beetles as forensic indicators. Insects. (2024) 15:711. doi: 10.3390/insects15090711
11. Michalski M and Nadolski J. Thermal conditions in selected urban and semi-natural habitats, important for the forensic entomology. Forensic Sci Int. (2018) 287:153–62. doi: 10.1016/j.forsciint.2018.03.042
12. Brown BV and Vendetti JE. Megaselia steptoeae (Diptera: Phoridae): specialists on smashed snails. Biodiversity Data J. (2020) 8:e50943. doi: 10.3897/BDJ.8.e50943
13. Grundmann B, Manko P, and Oboňa J. Overlooked insects in neglected ecosystem: new records of Phoridae for Slovakia discovered in rural environment. Hist naturalis bulgarica. (2024) 46:109–18. doi: 10.48027/hnb.46.043
14. Zuha RM, Jenarthanan LX, Disney RH, and Omar B. Multiple species of scuttle flies (Diptera: Phoridae) as contaminants in forensic entomology laboratory insect colony. Trop BioMed. (2015) 32:568–72.
15. Lutz L, Zehner R, Verhoff MA, Bratzke H, and Amendt J. It is all about the insects: a retrospective on 20 years of forensic entomology highlights the importance of insects in legal investigations. Int J Legal Med. (2021) 6:2637–2651. doi: 10.1007/s00414-021-02628-6
16. Sukontason K, Sukontason K, Vichairat K, Piangjai S, Lertthamnongtham S, Vogtsberger RC, et al. The first documented forensic entomology case in Thailand. J Med Entomology. (2001) 38:746–8. doi: 10.1603/0022-2585-38.5.746
17. Hu G, Wang M, Wang Y, Liao M, Hu J, Zhang Y, et al. Estimation of post-mortem interval based on insect species present on a corpse found in a suitcase. Forensic Sci Int. (2020) 306:110046. doi: 10.1016/j.forsciint.2019.110046
18. Reibe S and Madea B. Use of Megaselia scalaris (Diptera: Phoridae) for post-mortem interval estimation indoors. Parasitol Res. (2010) 106:637–40. doi: 10.1007/s00436-009-1713-5
19. Bugelli V, Forni D, Bassi LA, Di Paolo M, Marra D, Lenzi S, et al. Forensic entomology and the estimation of the minimum time since death in indoor cases. J Forensic Sci. (2014) 60:525–31. doi: 10.1111/1556-4029.12647
20. Syamsa R, Omar B, Zuha R, Faridah M, Swarhib M, Hidayatulfathi O, et al. Forensic entomology of high-rise buildings in Malaysia: Three case reports. Trop Biomed. (2015) 32:291–299.
21. Thevan K, Disney RHLH, and Ahmad AH. First records of two species of Oriental scuttle flies (Diptera: Phoridae) from forensic cases. Forensic Sci Int. (2010) 195:e5–7. doi: 10.1016/j.forsciint.2009.10.020
22. Li L, Wang Y, Liao M, Zhang Y, Kang C, Hu G, et al. The postmortem interval of two decedents and two dog carcasses at the same scene based on forensic entomology. Insects. (2022) 13:215. doi: 10.3390/insects13020215
23. Ivorra T, Rahimi R, Zaini NA, and Heo CC. Reporting beetles in a mummified human corpse in Malaysia: Necrobia rufipes (Fabricius 1781) (Coleoptera: Cleridae) and Dermestes maculatus De Geer 1774 (Coleoptera: Dermestidae). Legal Med. (2023) 62. doi: 10.1016/j.legalmed.2023.102238
24. Disney R and Manlove J. First occurrences of the Phorid, Megaselia abdita, in forensic cases in Britain. Med Veterinary Entomology. (2005) 19:489–91. doi: 10.1111/j.1365-2915.2005.00593.x
25. Manlove JD and Disney RHL. The use of Megaselia abdita (Diptera: Phoridae) in forensic entomology. Forensic Sci Int. (2008) 175:83–4. doi: 10.1016/j.forsciint.2007.08.001
26. Disney RHL and Manlove JD. First report of Triphleba nudipalpis (Becker) (Diptera: Phoridae) in a forensic case. Forensic Sci Int. (2009) 191:e1–3. doi: 10.1016/j.forsciint.2009.07.006
27. García-Rojo AM, Martínez-Sánchez A, López R, García de la Vega JM, Rica M, González M, et al. A mathematical model applied for assisting the estimation of PMI in a case of forensic importance. First record of Conicera similis (Diptera: Phoridae) in a corpse. Forensic Sci Int. (2013) 231:e11–8. doi: 10.1016/j.forsciint.2013.05.038
28. Merritt RW, Snider R, De Jong JL, Benbow ME, Kimbirauskas RK, and Kolar RE. Collembola of the grave: a cold case history involving arthropods 28 years after death. J Forensic Sci. (2007) 52:1359–61. doi: 10.1111/j.1556-4029.2007.00568.x
29. Ivorra T, Martínez-Sánchez A, Velásquez Y, Disney RHL, and Rojo S. Estimation of PMI for human remains wrapped in a blanket, with first report of Puliciphora rufipes (Phoridae) and Synthesiomyia nudiseta (Muscidae) outdoors in SE Spain. Legal Med. (2022) 57:102077. doi: 10.1016/j.legalmed.2022.102077
30. Amendt J, Goff ML, Campobasso CP, and Grassberger M. Current concepts in forensic entomology. Dordrecht, London: Springer (2010).
31. Pastula EC and Merritt RW. Insect arrival pattern and succession on buried carrion in michigan. J Med Entomology. (2013) 50:432–9. doi: 10.1603/me12138
32. Al-Zahrani O, Al-Khalifa MS, Al-Qahtni AH, and Al-Mekhlafi FA. Decomposition and dipteran succession on buried rabbits carcasses. Saudi J Biol Sci. (2023) 30:103822. doi: 10.1016/j.sjbs.2023.103822
33. Villet M. Borror and Delong’s Introduction to the Study of Insects, CA Triplehorn & NF Johnson: book review. Afr Entomology. (2005) 13:393–4.
34. Rodriguez WC and Bass WM. Decomposition of buried bodies and methods that may aid in their location. J Forensic Sci. (1985) 30:836–52. doi: 10.1520/JFS11017J
35. Alfsdotter C and Petaros A. Outdoor human decomposition in Sweden: A retrospective quantitative study of forensic-taphonomic changes and postmortem interval in terrestrial and aquatic settings. J forensic Sci. (2021) 66:1348–63. doi: 10.1111/1556-4029.14719
36. Simmons T, Cross PA, Adlam RE, and Moffatt C. The influence of insects on decomposition rate in buried and surface remains. J Forensic Sci. (2010) 55:889–92. doi: 10.1111/j.1556-4029.2010.01402.x
37. Guo Y, Li L, Liao M, Wang J, and Wang Y. Thick quilt may severely impact the estimation of postmortem interval using forensic entomology-based methods -two case reports. J forensic legal Med. (2023) 95:102501. doi: 10.1016/j.jflm.2023.102501
38. Pereira AJ, Breglia GA, and Uzal MH. Forensic entomology in homicide cases: study of a corpse found inside a buried vehicle. Forensic science medicine pathology. (2024) 21:358–365. doi: 10.1007/s12024-024-00889-2
39. Brownlow L, Young S, Fernández-Grandon M, and Hopkins RJ. Case closed - Wrappings and encasement delays and reduces fly presence on body parts. Forensic Sci Int. (2023) 342:111542. doi: 10.1016/j.forsciint.2022.111542
40. Ahmad A, Ahmad AH, Dieng H, Satho T, Ahmad H, Aziz AT, et al. Cadaver wrapping and arrival performance of adult flies in an oil palm plantation in northern peninsular Malaysia. J Med Entomology. (2011) 48:1236–46. doi: 10.1603/me10247
41. Whitaker A. Development of blowflies (Diptera: Calliphoridae) on pig and human cadavers: implications for forensic entomology casework. King’s College, London,UK (2014). Doctoral Thesis.
42. Zuha RM, See HW, Disney RHL, and Omar B. First record of genus Puliciphora Dahl (Diptera: Phoridae) associated with rabbit carcasses placed in concealed environments in Malaysia. Forensic Sci Int. (2014) 245:e36–7. doi: 10.1016/j.forsciint.2014.10.034
43. Raja MZ, See H-W, Disney RHL, and Baharudin O. Scuttle flies (Diptera: phoridae) inhabiting rabbit carcasses confined to plastic waste bins in Malaysia include new records and an undescribed species. Trop Life Sci Res. (2017) 28:131–43. doi: 10.21315/tlsr2017.28.1.9
44. Permadi AA, Al-Asytar AM, Chandra IJ, Augustin S, Risriani VM, Virnia, et al. A mathematical model for estimating post-mortem interval on wrapped and unwrapped carcasses. In: AIP conference proceedings. Melville: AIP Publishing LLC (2022). p. 020012.
45. Teo CH, Hing HL, Hamzah NH, and Hamzah SPAA. The effect of different coverings on total body score development of buried carcasses. Malaysian J Med sciences: MJMS. (2021) 28:103–12. doi: 10.21315/mjms2021.28.4.11
46. Taleb M, Tail G, Djedouani B, and Akgz HN. Impact of plastic wrapping on carcass decomposition and arthropod colonisation in northern Africa during spring. Sci justice: J Forensic Sci Soc. (2022) 62:117–27. doi: 10.1016/j.scijus.2021.11.005
47. Weger M, Cross P, and Cunliffe R. The effect of plastic bag containment of the head on the rate and pattern of decomposition. J Forensic Leg Med. (2020) 71:101922. doi: 10.1016/j.jflm.2020.101922
48. Bu H, Hu J, Han F, Wang L, Chen Q, Cheng P, et al. Polypropylene/Poly(butylene adipate-co-terephthalate) Breathing Film for Inhibiting Pseudomonas and Maintaining Microbial Communities and Postharvest Quality of Allium mongolicum Regel during Storage. Foods. (2023) 12. doi: 10.3390/foods12183370
49. Jaworski T and Hilszczański J. The effect of temperature and humidity changes on insects development their impact on forest ecosystems in the context of expected climate change. Leśne Prace Badawcze (Forest Research Papers) (2013) 74:345–355. doi: 10.2478/frp-2013-0033
50. Notter SJ and Stuart BH. The effect of body coverings on the formation of adipocere in an aqueous environment. J forensic Sci. (2012) 57:120–5. doi: 10.1111/j.1556-4029.2011.01943.x
51. Hill MA and Pokines JT. Comparative analysis of fetal pig decomposition processes in burials of variable depths and wrapping. J Forensic Sci. (2022) 67:2192–202. doi: 10.1111/1556-4029.15120
52. Kim H, Shin SE, Ko KS, and Park SH. The application of mitochondrial COI gene-based molecular identification of forensically important scuttle flies (Diptera: phoridae) in korea. BioMed Res Int. (2020) 2020:6235848. doi: 10.1155/2020/6235848
53. Talebzadeh F, Ghadipasha M, Gharedaghi J, Yeksan N, Akbarzadeh K, and Oshaghi MA. Insect fauna of human cadavers in tehran district. J arthropod-borne Dis. (2017) 11:363–70.
54. HEO CCH, Latif B, Kurahashi H, Tan SH, Chew WK, Nazni WA, et al. Preliminary observations of necrophagous dipteran activities and a new host record of parasitoids on a high-rise building. Polish J Entomology. (2017) 86:165–80. doi: 10.1515/pjen-2017-0011
55. Oliveira DL and Vasconcelos SD. Vertical location of ephemeral resources by adult Diptera: Implications for the colonization of cadavers in high-rise buildings. Forensic Sci Int. (2021) 324:110827. doi: 10.1016/j.forsciint.2021.110827
56. Gudipati M and Jampanapalli SR. Oral myiasis in a 6-year-old child with cerebral palsy: A case report. Int J Clin Pediatr Dent. (2024) 17:1281–4. doi: 10.5005/jp-journals-10005-2983
57. Sharif S, Singh CP, Athar B, Khan MK, and Qamar A. Forensic and ecological significance of necrophagous insects: Insights from animal carcasses, human cadavers, and myiasis patients. Legal Med (Tokyo Japan). (2024) 71:102544. doi: 10.1016/j.legalmed.2024.102544
58. Komori K, Hara K, Smith KG, Oda T, and Karamine D. A case of lung myiasis caused by larvae of Megaselia spiracularis Schmitz (Diptera: Phoridae). Trans R Soc Trop Med Hygiene. (1978) 72:467–70. doi: 10.1016/0035-9203(78)90165-7
59. Vanin S, Mazzariol S, Menandro ML, Lafisca A, and Turchetto M. Myiasis by Megaselia scalaris (Diptera: Phoridae) in a python affected by pulmonitis. J Med entomology. (2013) 50:209–11. doi: 10.1603/me12165
60. Giangaspero A, Barlaam A, Pane S, Marchili MR, Muda AO, Putignani L, et al. Accidental nasal myiasis caused by megaselia rufipes (Diptera: phoridae) in a child. J Med entomology. (2021) 58:121–4. doi: 10.1093/jme/tjaa184
61. Bugelli V, Tarozzi I, Galante N, Bortolini S, and Franceschetti L. Review on forensic importance of myiasis: Focus on medicolegal issues on post-mortem interval estimation and neglect evaluation. Legal Med (Tokyo Japan). (2023) 63:102263. doi: 10.1016/j.legalmed.2023.102263
62. Durão C, Barros A, and Campos P. A rare case of digital myiasis. J infection Public Health. (2017) 10:886–7. doi: 10.1016/j.jiph.2016.11.002
63. Bansode S and More V. Effects of temperature variation on the life cycle of the forensically important Calliphoridae fly Chrysomya rufifacies (Macquart 1843) (Diptera). Int J Trop Insect Sci. (2024) 44:747–57. doi: 10.1007/s42690-024-01189-6
64. Wang Y, Zhang Y, Hu G, Wang M, Zhu R, Zhai Y, et al. Development of Megaselia spiracularis (Diptera: Phoridae) at different constant temperatures. J thermal Biol. (2020) 93:102722. doi: 10.1016/j.jtherbio.2020.102722
65. Thomas JK, Sanford MR, Longnecker M, and Tomberlin JK. Effects of temperature and tissue type on the development of megaselia scalaris (Diptera: phoridae). J Med entomology. (2016) 53:519–25. doi: 10.1093/jme/tjw019
66. Zuha RM and Omar B. Developmental rate, size, and sexual dimorphism of Megaselia scalaris (Loew) (Diptera: Phoridae): its possible implications in forensic entomology. Parasitol Res. (2014) 113:2285–94. doi: 10.1007/s00436-014-3883-z
67. Feng D-X, Wu J, and Sun D-P. Intrapuparial age estimation of forensically important dohrniphora cornuta (Diptera: phoridae). J Med entomology. (2021) 58:616–24. doi: 10.1093/jme/tjaa219
68. Shikano I, Woolcott J, Cloonan K, Andreadis S, and Jenkins NE. Biology of mushroom phorid flies, megaselia halterata (Diptera: phoridae): effects of temperature, humidity, crowding, and compost stage. Environ entomology. (2021) 50:149–53. doi: 10.1093/ee/nvaa142
69. Soszynska A. The influence of environmental factors on the supranivean activity of flies (Diptera) in Central Poland. Eur J Entomology. (2004) 101:481–90. doi: 10.14411/eje.2004.068
70. Soszyńska-Maj A, Paasivirta L, and Giłka W. Why on the snow? Winter emergence strategies of snow-active Chironomidae (Diptera) in Poland. Insect Sci. (2016) 23:754–70. doi: 10.1111/1744-7917.12223
71. Disney RHL. Duration of development of two species of carrionbreeding scuttle flies and forensic implicati. Med Vet Entomol. (2005) 19:229–235. doi: 10.1111/j.0269-283X.2005.00562.x
72. Oliveira DL, Soares TF, and Vasconcelos SD. Effect of bait decomposition on the attractiveness to species of Diptera of veterinary and forensic importance in a rainforest fragment in Brazil. Parasitol Res. (2016) 115:449–55. doi: 10.1007/s00436-015-4811-6
73. Carter DO, Yellowlees D, and Tibbett M. Moisture can be the dominant environmental parameter governing cadaver decomposition in soil. Forensic Sci Int. (2010) 200:60–6. doi: 10.1016/j.forsciint.2010.03.031
74. Ong S-Q, Ahmad H, and Tan EH. Substrate moisture affects the development of megaselia scalaris (Diptera: phoridae): an implication of the growth circumstances of the fly in forensic entomology. Environ entomology. (2018) 47:1582–5. doi: 10.1093/ee/nvy127
75. Zou T-L, Feng D-X, Huang G-Y, Sun D-P, and Dai S-T. Species composition and succession of necrophagous insects on small buried baits in China. J Med Entomology. (2022) 59:1182–90. doi: 10.1093/jme/tjac045
76. Anderson GS. Factors that influence insect succession on carrion. In: Forensic entomology., 3rd edition ed, vol. 37. United Kingdom: CRC press (2019).
77. AL-Mekhlafi FA, Al-Zahrani O, Al-Qahtni AH, and Al-Khalifal MS. Decomposition of buried rabbits and pattern succession of insect arrival on buried carcasses. Int J Trop Insect Sci. (2024) 44:1037–1044. doi: 10.1007/s42690-024-01203-x
78. Iancu L, Junkins EN, and Purcarea C. Characterization and microbial analysis of first recorded observation of Conicera similis Haliday (Diptera: Phoridae) in forensic decomposition study in Romania. J Forensic Legal Med. (2018) 58:50–5. doi: 10.1016/j.jflm.2018.04.012
79. Han W, Feng D, and Tang Y. The Effect of Soil Type and Moisture on the Development of Forensically Important Megaselia scalaris and Dohrniphora cornuta (Diptera: Phoridae). Insects. (2024) 15:666. doi: 10.3390/insects15090666
80. Ren L, Shang Y, Chen W, Meng F, Cai J, Zhu G, et al. A brief review of forensically important flesh flies (Diptera: Sarcophagidae). Forensic Sci Res. (2018) 3:16–26. doi: 10.1080/20961790.2018.1432099
81. Chophi R, Sharma S, Sharma S, and Singh R. Forensic entomotoxicology: Current concepts, trends and challenges. J forensic legal Med. (2019) 67:28–36. doi: 10.1016/j.jflm.2019.07.010
82. Pounder DJ. Forensic entomo-toxicology. J - Forensic Sci Soc. (1991) 31:469–72. doi: 10.1016/s0015-7368(91)73189-7
83. Bugelli V, Papi L, Fornaro S, Stefanelli F, Chericoni S, Giusiani M, et al. Entomotoxicology in burnt bodies: a case of maternal filicide-suicide by fire. Int J legal Med. (2017) 131:1299–306. doi: 10.1007/s00414-017-1628-0
84. Miller ML, Lord WD, Goff ML, Donnelly B, Mcdonough ET, and Alexis JC. Isolation of amitriptyline and nortriptyline from fly puparia (phoridae) and beetle exuviae (dermestidae) associated with mummified human remains. J Forensicences. (1994) 39:1305–13. doi: 10.1520/JFS13717J
85. Li T, Yin Y, Zhang K, Li Y, Kong X, Liu D, et al. Ecotoxicity effect of aspirin on the larvae of Musca domestica through retinol metabolism. Ecotoxicology Environ Saf. (2024) 270:115845. doi: 10.1016/j.ecoenv.2023.115845
86. Quijano-Mateos A, Castillo-Alanis A, Pedraza-Lara CS, and Bravo-Gómez ME. Evaluation of the effect of clonazepam and its metabolites on the life cycle of Megaselia scalaris (Loew)(Diptera: Phoridae). Sci Justice. (2024) 64:460–5. doi: 10.1016/j.scijus.2024.07.002
87. Castillo-Alanis LA, González-Hernández AE, Quijano-Mateos A, Pedraza-Lara CS, Villavicencio-Queijeiro A, and Bravo-Gómez ME. Standardization of a culture medium for megaselia scalaris (Diptera: phoridae) for entomotoxicological studies. J Med Entomology. (2020) 57:1421–31. doi: 10.1093/jme/tjaa088
88. Bhardwaj T, Sharma S, Dalal J, and Verma K. The implication of morphometrics and growth rate of dipteran flies in forensic entomotoxicology research: a review. Die Naturwissenschaften. (2020) 107:50. doi: 10.1007/s00114-020-01707-9
89. Salimi M, Rassi Y, Chatrabgoun O, Kamali A, Oshaghi MA, Shiri-Ghaleh V, et al. Toxicological analysis of insects on the corpse: A valuable source of information in forensic investigations. J arthropod-borne Dis. (2018) 12:219–31. doi: 10.18502/jad.v12i3.74
90. Alcaine-Colet A, Wotton KR, and Jimenez-Guri E. Rearing the scuttle fly Megaselia scalaris (Diptera: Phoridae) on industrial compounds: implications on size and lifespan. PeerJ. (2015) 3:e1085. doi: 10.7717/peerj.1085
91. Jensen P and Trumble JT. Ecological consequences of bioavailability of metals and metalloids in insects. Recent Res Dev Entomol. (2003) 42:1–17.
92. Trumble JT and Jensen PD. Ovipositional response, developmental effects and toxicity of hexavalent chromium to Megaselia scalaris, a terrestrial detritivore. Arch Environ Contamination Toxicol. (2004) 46:372–6. doi: 10.1007/s00244-003-3007-8
93. Jensen PD, Rivas MD, and Trumble JT. Developmental responses of a terrestrial insect detritivore, Megaselia scalaris (Loew) to four selenium species. Ecotoxicology (London England). (2005) 14:313–22. doi: 10.1007/s10646-003-6368-x
94. Castillo-Alanis LA, Lira-Paredes AA, Pedraza-Lara C, Quijano-Mateos A, and Bravo-Gómez ME. Effect of malathion on the development of megaselia scalaris (Loew 1866) (Diptera: phoridae), a forensically important fly. J Med entomology. (2022) 59:826–34. doi: 10.1093/jme/tjac025
95. Rumiza AR, Khairul O, Zuha RM, and Heo CC. An observation on the decomposition process of gasoline-ingested monkey carcasses in a secondary forest in Malaysia. Trop biomedicine. (2010) 27:373–83.
96. Zulaikha ABS and Zuha RM. Nocturnal oviposition of the forensic scuttle fly, Megaselia scalaris (Loew) (Diptera: Phoridae) indoors. Egyptian J Forensic Sci. (2016) 6:489–91. doi: 10.1016/j.ejfs.2016.02.002
97. Bostock E, Green EW, Kyriacou CP, and Vanin S. Chronobiological studies on body search, oviposition and emergence of Megaselia scalaris (Diptera, Phoridae) in controlled conditions. Forensic Sci Int. (2017) 275:155–9. doi: 10.1016/j.forsciint.2017.03.002
98. Wang Y, Gong Q, Liu ZJ, Wang M, Xu W, Wang YH, et al. Research progress on developmental biology of sarcosaprophagous insects. Fa Yi Xue Za Zhi. (2021) 37:673–82. doi: 10.12116/j.issn.1004-5619.2020.401216
99. Cui B. Research progress on the wing and veins of Diptera Phoridae. Int J Biol. (2010) 2:210. doi: 10.5539/ijb.v2n2p210
100. Pallavi J, Snehal H, Kolipakala RS, Salazar D, Hanbar M, Chiramel LB, et al. A complete morphological characterization of all life stages of the phorid fly Megaselia scalaris. Sci Rep. (2023) 13. doi: 10.1038/s41598-023-50200-6
101. Disney R. Natural history of the scuttle fly, Megaselia scalaris. Annu Rev Entomol. (2008) 53:39–60. doi: 10.1146/annurev.ento.53.103106.093415
102. Harrison DA and Cooper RL. Characterization of development, behavior and neuromuscular physiology in the phorid fly, Megaselia scalaris. Comp Biochem Physiol Part A: Mol Integr Physiol. (2003) 136:427–39. doi: 10.1016/S1095-6433(03)00200-9
103. Binsha P, Thomas SK, and Sunny J. Biological Control Potential of the Scuttle Fly, Megaselia scalaris (Loew)(Diptera: Phoridae), on the Home-Invading Pest Luprops tristis (Fabricius)(Coleoptera: Tenebrionidae). Coleopterists Bull. (2020) 74:331–6. doi: 10.1649/0010-065X-74.2.331
104. Szleszkowski Ł., Kadej M, Ogórek R, Thannhäuser A, Dobrowolski MA, and Jurek T. Salt mummification” - atypical method of embalming a corpse. Int J Legal Med. (2022) 136:1829–40. doi: 10.1007/s00414-022-02855-5
105. Campobasso CP, Henry R, Disney L, and Introna F. A case of Megaselia scalaris (Loew)(Dipt., Phoridae) breeding in a human corpse. Anil Aggrawal’s Internet J Forensic Med Toxicol. (2004) 5:3–5.
106. Greenberg B and Wells JD. Forensic use of Megaselia abdita and M. scalaris (Phoridae: Diptera): case studies, development rates, and egg structure. J Med Entomology. (1998) 35:205–9. doi: 10.1093/jmedent/35.3.205
107. Grassberger M and Frank C. Initial study of arthropod succession on pig carrion in a central European urban habitat. J Med Entomology. (2004) 41:511–23. doi: 10.1603/0022-2585-41.3.511
108. Zhang Y, Li L, Liao M, Kang C, Hu G, Guo Y, et al. Development of Megaselia scalaris at constant temperatures and its significance in estimating the time of death. Int J Legal Med. (2024) 138:97–106. doi: 10.1007/s00414-023-02993-4
109. Elsheakh MH, Mohamed HA, Abied MK, and Ebrahim AM. Survey and taxonomy of phorid flies (Phoridae: Diptera) with newly recorded genus in Egypt. Egyptian J Plant Prot Res Institute. (2024) 7:19–30. doi: 10.4314/ejppri.v7i1
110. Kumara T, Disney R, Hassan AA, Flores M, Hwa TS, Mohamed Z, et al. Occurrence of oriental flies associated with indoor and outdoor human remains in the tropical climate of north Malaysia. J Vector Ecol. (2012) 37:62–8. doi: 10.1111/j.1948-7134.2012.00200.x
111. Feng D-X and Liu G-C. Morphology of immature stages of Dohrniphora cornuta (Bigot) (Diptera: Phoridae). Microscopy Res technique. (2012) 75:1528–33. doi: 10.1002/jemt.22096
112. Disney RHL, Garcia-Rojo A, Lindström A, and Manlove JD. Further occurrences of Dohrniphora cornuta (Bigot) (Diptera, Phoridae) in forensic cases indicate likely importance of this species in future cases. Forensic Sci Int. (2014) 241:e20–22. doi: 10.1016/j.forsciint.2014.05.010
113. Velásquez Y, Magaña C, Martínez-Sánchez A, and Rojo S. Diptera of forensic importance in the Iberian Peninsula: larval identification key. Med veterinary entomology. (2010) 24:293–308. doi: 10.1111/j.1365-2915.2010.00879.x
114. Bourel B, Tournel G, Hédouin V, and Gosset D. Entomofauna of buried bodies in northern France. Int J Legal Med. (2004) 118:215–20. doi: 10.1007/s00414-004-0449-0
115. Disney R, Henderson I, Perry J, and Clements R. Phoridae (Diptera) from English pasture soils. Pedobiologia. (1981) 22:366–78. doi: 10.1016/S0031-4056(23)03701-0
116. Disney R. Scuttle flies (Diptera: Phoridae) as parasites of earthworms (Oligochaeta: Lumbricidae). Br J entomology Natural history. (1991) 4:11–3.
117. Lundt H. Ecological observations about the invasion of insects into carcasses buried in soil. Pedobiologia. (1964) 4:158–80. doi: 10.1016/S0031-4056(22)00150-0
119. Henry R and Disney L. Seven new species of New World Puliciphora Dahl (Diptera: Phoridae) with a new key to the Neotropical species. Zootaxa. (2003) 162:1–22-21–22. doi: 10.11646/zootaxa.162.1.1
120. Sinclair BJ and Disney RHL. Some scuttle flies (Diptera: phoridae) of the galápagos islands. Tijdschrift Voor Entomologie. (2008) 151:115–32. doi: 10.1163/22119434-900000257
121. Dai S-T, Feng D-X, and Sun D-P. Characterization and Phylogenetic Analysis of the Complete Mitochondrial Genomes of Two Tiny Necrophagous Phorid Flies, Metopina sagittata and Puliciphora borinquenensis (Diptera: Phoridae). J Med entomology. (2022) 59:120–8. doi: 10.1093/jme/tjab152
122. Chakraborty A, Naskar A, Parui P, and Banerjee D. Developmental variation of Indian thermophilic variety of scuttle fly megaselia (Megaselia) scalaris (Loew 1866) (Diptera: phoridae) on different substrates. Scientifica. (2016) 2016:4257081. doi: 10.1155/2016/4257081
123. Burcham ZM, Belk AD, McGivern BB, Bouslimani A, Ghadermazi P, Martino C, et al. A conserved interdomain microbial network underpins cadaver decomposition despite environmental variables. Nat Microbiol. (2024) 9:33. doi: 10.1038/s41564-023-01580-y
124. Holman AP, Pickett DN, West H, Tarone AM, and Kurouski D. Portable Fourier-transform infrared spectroscopy and machine learning for sex determination in third instar Chrysomya rufifacies larvae. J Forensic Sci. (2025) 70:1468–79. doi: 10.1111/1556-4029.70054
Keywords: corpses, forensic entomology, insect, phorid fly, postmortem interval
Citation: Guo Y, Luo Y, Ma Y, Luciano A, Huang J, Li Y, Wang S, Niu Y, Li Z, Cai J and Meng F (2026) Wings on concealed corpse: the forensic importance of scuttle flies (Diptera: Phoridae). Front. Insect Sci. 5:1727384. doi: 10.3389/finsc.2025.1727384
Received: 17 October 2025; Accepted: 12 December 2025; Revised: 28 November 2025;
Published: 19 January 2026.
Edited by:
Atilano Contreras-Ramos, National Autonomous University of Mexico, MexicoReviewed by:
Humberto Quiroz-Martinez, Autonomous University of Nuevo León, MexicoCopyright © 2026 Guo, Luo, Ma, Luciano, Huang, Li, Wang, Niu, Li, Cai and Meng. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Fanming Meng, bWVuZ2Zhbm1pbmcxOTg0QDE2My5jb20=; Jifeng Cai, Y2pmX2ppZmVuZ0AxNjMuY29t
†These authors share first authorship