Carcases and mites

Exp Appl Acarol (2009) 49:45–84 DOI 10.1007/s10493-009-9287-6 Carcases and mites Henk R. Braig Æ M. Alejandra Perotti Received: 5 June 2009 / Accepted: 16 June 2009 / Published online: 24 July 2009 Ó Springer Science+Business Media B.V. 2009 Abstract Mites are involved in the decomposition of animal carcases and human corpses at every stage. From initial decay at the fresh stage until dry decomposition at the skeletal stage, a huge diversity of Acari, including members of the Mesostigmata, Prostigmata, Astigmata, Endeostigmata, Oribatida and Ixodida, are an integral part of the constantly changing food webs on, in and beneath the carrion. During the desiccation stage in wave 6 of Mégnin’s system, mites can become the dominant fauna on the decomposing body. Under conditions unfavourable for the colonisation of insects, such as concealment, low temperature or mummification, mites might become the most important or even the only arthropods on a dead body. Some mite species will be represented by a few specimens, whereas others might build up in numbers to several million individuals. Astigmata are most prominent in numbers and Mesostigmata in diversity. More than 100 mite species and over 60 mite families were collected from animal carcases, and around 75 species and over 20 families from human corpses. Keywords Carrion
Carcass
Corpse
Cadaver
Animal decomposition
Necrophagy
Necrophagia
Succession
Post mortem interval Introduction Corpses of humans and carcases of animals represent biocenoses that are often composed of complicated food webs. Especially under the combined influence of residential bacteria from the gut and introduced blow or flesh flies, the decomposition of a recently deceased body can proceed very rapidly, resulting in a constantly changing habitat for necrophilous and necrophagous arthropods and other animals and fungi. These changes might be H. R. Braig (&) School of Biological Sciences, Bangor University, Deiniol Road, Bangor, Wales LL57 2UW, UK e-mail: h.braig@bangor.ac.uk M. A. Perotti School of Biological Sciences, University of Reading, Whiteknights, Reading, Berkshire RG6 6AS, UK 123 46 Exp Appl Acarol (2009) 49:45–84 considered as a succession of microhabitats or seral sequences, microseres, which might be divided into a series of definable stages that might be called microseral stages. Insect species dominate the serially changing populations on carcases. However, mites are receiving increased recognition as a part of forensic biology (Frost et al. 2009; Perotti and Braig 2009a; Perotti et al. 2009b). Mites are also involved in most stages of decomposition of animal and human remains. This paper tries to list the most abundant mite fauna associated with decomposition. Waves of arthropods Early work on decomposition in forensic medicine was inspired by case observations of the arthropod fauna associated with exposed human corpses. Jean Pierre Mégnin in Paris, France, organised his observations in his book La Faune des Cadavres [The Fauna of Carcases], where he observed that arthropods appear in 8 distinct waves on the carcases of humans. He illustrated this with 19 forensic case studies described in detail (Mégnin 1894). A short summary of the 8 waves was published a year later (Mégnin 1895). There remains an oddity in Mégnin’s legacy. Specimens of the corpse fly Hydrotaea capensis recovered from 1 year-old corpses from the cemetery of Saint Nazaire in Paris were assigned by Mégnin to wave 5 and to an otherwise unknown wave 9 (Pont and Matile 1980). Over time, several more insect species have been added to the list of waves of arthropods (Table 1). In Mégnin’s original observations, an entire wave, the sixth, was composed of only mites. Later on, Leclercq added mites also to the very first wave (Leclercq and Verstraeten 1993). Several other authors have added additional species to the list of waves. Porta in Parma, Italy, distinguished 9 waves of arthropods associated with ten stages of human decomposition. In his system, waves 6 and 7 were, among others, characterised by larvae, nymphs and adults of Acari. These 2 waves represent the initial and final pre-skeletal stages, each lasting for 3–4 months for exposed and for concealed corpses (Porta 1929). At the skeletal stage, only small numbers of adult mites were recovered by Porta. Mégnin’s appreciation of mites in a forensic context has been acknowledged early on by forensic entomologists and pathologists (Graells 1886; Rı́os 1902a, b; Lecha-Marzo 1917; Porta 1929). However, the proposed succession of insects and Mégnin’s interpretations were questioned over time by many (Strauch 1912; Wyss and Cherix 2006). Mégnin’s work on the arthropod succession on human corpses led him to describe several new species of mites and flies. Some of the species descriptions in La Faune des Cadavres are very brief and the associated drawings not particularly detailed. This has not been a problem in cases where subsequent workers have acknowledged Mégnin’s species descriptions and included them in their revisions. Serrator amphibius Mégnin (1894) is a revision by Mégnin himself of Tyroglyphus rostro-serratus Mégnin 1873 and should now be recognised as Histiostoma feroniarum (Dufour 1839) (Histiostomatidae, Astigmata). The identification of Serrator necrophagus Mégnin (1894) is more of a problem. Should it be considered as Histiostoma necrophagus (=? necrophori Dujardin) (Leclercq and Verstraeten 1988b)? According to OConnor (pers. comm.), S. necrophagus is a composite of Histiostoma and Myianoetus and as such unrecognisable. The two species Uropoda nummularia Mégnin (1894) (? Uropodidae Kramer 1881, Mesostigmata) and Trachynotus cadaverinus Mégnin (1894) (? Trachyuropodidae Berlese 1917, Mesostigmata) had not been taken up by a systematic acarologist and their identity has remained a puzzle for a long time. Few authors have reproduced the characteristics of 123 Exp Appl Acarol (2009) 49:45–84 47 Table 1 Based on 15 years of experience at the Paris morgue, Mégnin described 8 waves, squads or periods of arthropod succession on human corpses exposed to the air (escouades or séries des travailleurs de la mort [sections or series of death workers or gravediggers of nature (Gaudry 2002)]) Faunal succession as established by Mégnin on exposed human corpses 1st wave – bodies fresh; normally, first 48 h but can last for 3 months after death Muscidae Musca domestica, house fly M. autumnalis (=M. corvina), face or autumn house fly Muscina stabulans (=Curtonevra stabulans), false stable fly Stratiomyidae Hermetia illucens, black soldier fly Phoridae humpbacked or scuttle flies Calliphoridae Calliphora vomitoria, holarctic blue blow fly C. vicina (=C. erythrocephala), European bluebottle fly Chrysomya albiceps, blow fly Lucilia spp., greenbottle flies Protophormia terraenovae, bird’s nest screwworm fly Phormia regina, black blow fly Acari mites 2nd wave – decomposition commenced, odour developing; 48–72 h but can last for the first 3 months after death Muscidae Hydrotaea dentipes, sweat fly Calliphoridae Lucilia caesar, golden greenbottle fly Lucilia sericata (=Phaenicia sericata), sheep blow fly Cynomya mortuorum, bluebottle fly Sarcophagidae Sarcophaga carnaria, grey flesh fly S. arvensis, flesh fly S. laticrus (=Myophora laticrus), flesh fly S. (Liopygia) argyrostoma (=Parasarcophaga argyrostoma), flesh fly Staphylinidae Omalium rivulare, rove beetle 3rd wave – fats becoming racid, butyric fermentation; 3–6 months after death Dermestidae Dermestes lardarius, larder or bacon beetle D. frischi, common hide beetle D. undulatus, skin beetle Pyralidae Aglossa pinguinalis, grease moth A. caprealis, fungus or murky meal moth 4th wave – caseous fermentation; 3–4 to 6–8 months after death Piophilidae Piophila casei, cheese skipper, jumping maggot P. petasionis, ham and cheese fly 123 48 Exp Appl Acarol (2009) 49:45–84 Table 1 continued Faunal succession as established by Mégnin on exposed human corpses Anthomyiidae Chortophila vicina (=Anthomyia vicina), banded fly Anthomyia pluvialis, banded fly A. vesicularis, banded fly Cleridae Korynetes caeruleus (=Corynetes violaceus), bone beetle K. ruficornis (=Corynetes coeruleus), blue hide beetle Necrobia ruficollis (=Corynetes ruficollis), red-shouldered ham beetle N. rufipes (=Corynetes rufipes), red-legged ham beetle N. violacea, black-legged ham beetle, blue corynetes Staphylinidae Omalium rivulare, rove beetle Fanniidae Fannia scalaris (=Anthomyia scalaris), latrine fly Milichiidae Madiza glabra, insect jackal Syrphidae Eristalis tenax, drone fly, rat-tailed maggot Brachyopa spp., hover flies Ephydridae Scatella fusca (=Teichomyza fusca), urine or urinal fly Heleomyzidae Tephrochlamys rufiventris, sun fly Drosophilidae vinegar flies Sciaridae dark-winged fungus gnats Sepsidae black scavenger flies Sphaeroceridae small dung flies Trichoceridae winter crane flies 5th wave – ammoniacal fermentation, black liquefaction, evaporation of sanious fluids; 4–5 to 8–9 months after death Piophilidae Thyreophora cynophila, skipper fly, considered extinct Centrophlebomyia anthropophaga (=Thyreophora anthropophaga), bone skipper, almost extinct C. furcata, bone skipper Dasyphlebomyia stylata, skipper fly Lonchaeidae Lonchaea nigrimana, lance fly Muscidae Hydrotaea capensis (=Ophyra cadaverina Mégnin,=Ophyra anthrax), dung or corpse fly H. leucostoma (=Ophyra leucostoma), black garbage or dump fly Phoridae Phora aterrima, scuttle fly Triphleba spp., humpbacked flies Silphidae Nicrophorus interruptus (=Necrophorus fossor), burying beetle 123 Exp Appl Acarol (2009) 49:45–84 49 Table 1 continued Faunal succession as established by Mégnin on exposed human corpses N. humator, black sexton beetle N. investigator, banded sexton beetle Necrodes littoralis (=Silpha littoralis), bent-legged silpha, shore sexton beetle Oiceoptoma noveboracensis (=Silpha noveboracensis), small or margined carrion beetle Silpha obscura, carrion beetle Histeridae Margarinotus brunneus (=Hister cadaverinus, H. impressus), clown beetle Gnathoncus rotundatus (=Saprinus rotondatus), carrion beetle Euspilotus assimilis (=Saprinus assimilis), clown beetle Saprinus semistriatus, striped clown beetle Hister foedatus, hister beetle Leiodidae Catops spp., round fungus beetle Nitidulidae Carpophilus spp., dried fruit beetles 6th wave – desiccation; 5–6 to 10–12 months after death Mesostigmata Dinychidae (Uropodidae) Leiodinychus krameri (=Uropoda nummularia Mégnin) ? Trachytidae Uroseius acuminatus (=Trachynotus cadaverinus Mégnin) ? Astigmata Acaridae Acarus siro (=Tyroglyphus siro, Tyrolichus casei) Tyrophagus longior (=Tyroglyphus longior, Tyroglyphus infestans) Histiostomatidae Histiostoma feroniarum (=Serrator amphibius Mégnin, Tyroglyphus rostro-serratus Mégnin) Serrator necrophagus Mégnin ? Glycyphagidae Glycyphagus destructor (=Glyciphagus cursor Mégnin, Glyciphagus spinipes) 7th wave – complete desiccation; after 8 months or 1–3 years after death Pyralidae Aglossa caprealis, fungus or murky meal moth Tineidae Tineola bisselliella, webbing clothes or carpet moth Tinea pellionella, case-making clothes moth Monopis laevigella (=M. rusticella), fur moth Dermestidae Attagenus pellio, fur beetle Anthrenus museorum, museum beetle Dermestes maculatus, leather, hide or bacon beetle Nitidulidae Omosita colon, pollen or sap beetle Trogidae Trox unistriatus, skin beetle 123 50 Exp Appl Acarol (2009) 49:45–84 Table 1 continued Faunal succession as established by Mégnin on exposed human corpses 8th wave – debris; over 3 years after death Tenebrionidae Tenebrio molitor, yellow mealworm beetle T. obscurus, dark mealworm beetle Anobiidae Ptinus brunneus, brown spider beetle Species aligned to the left in the list represent the species originally identified by Mégnin (1894), species more to the right are additions made by subsequent workers (Johnston and Villeneuve 1897; Leclerq 1969; Smith 1973, 1986; Leclercq and Verstraeten 1993; Gaudry 2002). For some of the additional species, the assignment of a species to a particular wave varies with the locality and author. The systematics of species has been adapted to current use; the original and one of its synonyms, where appropriate, are in parentheses. Species names with ‘?’ are discussed in the text. Where available, the vernacular name of the insect species is given, otherwise one of the common names of its family is used Mégnin’s species and often not in easily accessible publications, which might have contributed to them being overlooked (Rı́os 1902b; Porta 1929). In addition, the mite name T. cadaverinus is sometimes confused with a beetle species. However, these species have finally been identified as quite common and widespread mites. Athias-Binche (1994) recognises U. nummularia as a synonym of the round grain or round brown mite, Leiodinychus krameri (G & R Canestrini 1882) (Dinychidae or Uropodidae) and T. cadaverinus as Uroseius acuminatus (CL Koch 1847) (Trachytidae), which can be phoretic on the phorid fly Aphiochaeta rufipes. Mégnin differentiates between Glyciphagus spinipes Ch. Rob. and Glyciphagus cursor Mégnin (1894), both are now considered synonyms of the pilous or groceries mite Glycyphagus (Lepidoglyphus) destructor (Schrank 1781) (Glycyphagidae, Astigmata). Mégnin also differentiates between Tyroglyphus longior Gervais 1844 (Mégnin 1894) and Tyroglyphus infestans Berlese 1884 (Mégnin 1898), both are now synonyms of the seed mite Tyrophagus longior (Gervais 1844). However, the Tyrophagus species reported by Mégnin might have been a mixture of species (Perotti 2009). The forensically important bulb mite species Cœpophagus echinopus depictured in detail in Mégnin’s La Faune des Cadavres in 1894 is now recognised as Rhizoglyphus echinopus (Fumouze and Robin 1868) (Acaridae, Astigmata). All species in the genus Caloglyphus Berlese 1923 will be listed as Sancassania Oudemans 1916 (Acaridae, Astigmata) (Samšiňák 1960). Tyroglyphus mycophagus Mégnin 1874 became Caloglyphus mycophagus and is now S. berlesei (Michael 1903). Some consider it one species, according to Hughes and Baker these are two species, and Moniez in 1892 has described a mite species as Tyroglyphus mycophagus that is now recognised as S. chelone Oudemans 1916. In the early Spanish literature, mites of the genus Carpoglyphus (Carpoglyphidae, Astigmata) are listed as part of Mégnin’s mite-rich sixth wave but have not been reported since then (Lecha-Marzo 1917). The carrion or grave fly, Ophyra cadaverina Mégnin (1894) (Muscidae, Diptera), fifth wave, had been ignored by entomologists for some time. Around 85 years after the original publication in Mégnin’s book, a bottle was discovered by accident in the Natural History Museum in Paris with insects collected from corpses and labelled ‘Travailleurs de la Mort’. 123 Exp Appl Acarol (2009) 49:45–84 51 The bottle also contained three specimens of O. cadaverina that allowed the identification of Mégnin’s species as a junior synonym of O. capensis (Wiedemann, 1818; Pont and Matile 1980). Species in the genus Ophyria have meanwhile been transferred to the genus Hydrotaea, however, molecular studies place Ophyria species in a clade separate from Hydrotaea (Schnell e Schühli et al. 2004, 2007). The bottle must have been part of original material offered to the museum by Mégnin. Acarologists have not yet investigated whether some of the mites have been saved as well. It is surprising that Mégnin didn’t observe any mite species in wave 7, complete desiccation. The beetle species in this wave, Dermestes spp., Trox spp. and similar species, are well known for the large numbers and diversity of phoretic mites they carry (Perotti and Braig 2009b). Some taxa such as the grease and fungi moths, may appear subsequently in 2 separate waves; first with wave 3, when the body fats started oxidising, particularly Aglossa pinguinalis, and later with wave 7, when the carcase has dried out, mostly A. cuprealis. The species composition of insects and mites will vary with the region, temperature, season, amount of light and shade, level of concealment, presence of vertebrate scavengers and other environmental peculiarities. Interestingly, the species composition might even change with time. For example, several species of bone skippers, Thyreophora species, are so specialised to later stages of the decomposition of large carcases that they have become extinct or are close to extinction. Decomposing bone marrow may be the preferred larval diet or the protection provided by large bones might be essential for the survival of the larvae. These species only remain in small pockets in countries like India (Kashmir) where their existence depends on the availability of later stages of decomposition of large animal carcases like horses (Michelsen 1983). One expects that Indian elephants might provide an even better habitat for these flies. Ironically, Thyreophora is not only a skipper fly genus threatened by extinction, it is also an extinct suborder of shield-bearing dinosaurs. During the time of Mégnin, sufficient numbers of large animals seem to have been allowed to decompose completely in nature to enable the species to survive. Through human intervention, most large animal carcases are now removed from the land before they reach advanced stages of decomposition. Changes in human behaviour influence which species participate in the decomposition process. The time line of the 8 waves seems to have changed as well. Leclercq observed that the scuttle flies, Phoridae, no longer appear in wave 5 around 4–8 months after death but might arrive as early as week 3 and might also be found very late until several years after death. The mites no longer colonise the carcase as a compact wave between 6 and 12 months but in the experience of Leclercq, mites will arrive much earlier and more likely in 4 specific waves dependent on the physical state of decomposition of the carcase. He differentiates between the following appearances of the carcase as specific habitats for mites: ‘outright liquid [franchement aquatiques]’, ‘semi liquid [semi-aquatiques]’, ‘a little bit wet [peu hydrophiles]’ and ‘in the process of desiccation or dry [milieu en voie de dessication ou desséché]’ but didn’t assign specific species to each habitat (Leclercq and Verstraeten 1988a, 1993; Leclercq 2002). The waves of arthropods in Mégnin’s system overlap with each other; they often form a continuum where it becomes difficult to say where one particular wave ends and a subsequent wave starts. Environmental conditions like the degree of drying out of the carcase or the impact of vertebrate scavengers might prevent several waves of arthropods arriving at a carcase. Many insect species are habitat specific. Ants (Hymenoptera), not mentioned in Mégnin’s system, might be the numerically dominant species on a carcase under certain environmental conditions. And more critique has been expressed regarding individual waves and taxa. However, the acarological importance of this list is that most if not all of the insects arriving at the carcase might carry mites. Perhaps the easiest way to obtain a 123 52 Exp Appl Acarol (2009) 49:45–84 structured overview of the time line, the potential mite carriers and of the potential predators of mites still might be the use of Mégnin’s system. Stages of decomposition Currently the state of a carcase is described by a state of decomposition rather than by a wave of arthropod colonisation. Five stages (Table 2) are most commonly recognised for exposed and concealed carcases as described by Goff (2009). Six stages of decay are proposed for the decomposition of pig carcases in water (Payne and King 1972). Table 2 Terms of the most commonly recognised five stages of decomposition of vertebrate animals and humans 1 Initial decay, fresh stage Carcase appears fresh externally but is decomposing internally due to the activities of bacteria, protozoa and nematodes present in the animal before death. This stage begins at the moment of death and ends when bloating is first evident. The first organisms to arrive are blow flies and flesh flies. Eggs or larvae are deposited around the natural openings or wounds 2 Putrefaction, bloated stage Carcase swollen by gas produced internally, accompanied by odour of decaying flesh. Gasses produced by the metabolic activities of anaerobic bacteria first cause a slight inflation of the abdomen, and the corpse may later assume a fully inflated, balloon-like appearance. Internal carcase temperatures begin to rise as a combined result of putrefaction processes and metabolic heat of the fly larvae. Predatory taxa such as rove beetles arrive. Fluids seeping from natural body openings combined with ammonia produced by the fly larvae cause the soil beneath the carcase to become alkaline. Normal soil fauna will depart the area beneath the remains 3 Black putrefaction, active decay, decay stage Flesh of creamy consistency with exposed parts black. Body collapses as gases escape. Odour of decay very strong. The decay stage begins when the skin is broken, allowing gases to escape and the remains deflate. Diptera larvae from large feeding masses are the predominant taxa; Coleoptera arrive in numbers. Necrophagous and predatory taxa are observed in large numbers during the latter part of the stage. By the end of the stage, the blow and flesh flies will have departed the remains for pupariation. The fly larvae will have removed most of the flesh by the end 4 Butyric fermentation, advanced decay, post-decay stage Carcase drying out. Some flesh remains at first, and cheesy odour develops. Ventral surface of body mouldy from fermentation. Remains are reduced to skin, cartilage, and bones. Various beetle species will dominate and their diversity will increase; parasites and predators of beetles will increase as well. In wet habitats such as swamps and rain forests, beetles will be replaced by flies and other taxa 5 Dry decay, dry decomposition, skeletal stage, remains stage Carcase almost dry to complete dry; slow rate of decay. Only bones and hair remain. A gradual return of the normal soil fauna to the area beneath the remains. There is no definitive end point to this stage and some variations in the composition of the soil fauna may be detectable even years following the death depending on local conditions Some stages have (almost) interchangeable names given by different authorities, like butyric fermentation and advanced decay; others like butyric fermentation and post-decay overlap only partially. The last term at each stage is the one used by Goff (2009). The rough description of the stages follows Bornemissza (1957) for guinea pigs. The more detailed description follows Goff (1993) for pigs and humans 123 Exp Appl Acarol (2009) 49:45–84 53 Mites are numerous on carcases Mites are not a rarity on carcases. A few examples and citations from the literature might illustrate this. Acarina are numerous on pig carcases (Gill 2005). Butyric fermentation and advanced decay will attract mites in such numbers that they become visible to the naked eye. However, they are often mistaken for mould, which is present at that time as well, or for fine sawdust, as is emphasised by one of the classical chapters on forensic entomology (Haskell et al. 1997). Large quantities of mites give a fluffy appearance to decomposing pigs (Anderson et al. 2002). In a study of 43 dog carcases in Tennessee (USA), mites were sometimes distributed on the upper surface of carcases (Reed 1958). Where any skin was left by the skin feeders of the previous stage, an immense number of tyroglyphid mites consumed the remainder leaving nothing but bones of guinea pigs (Bornemissza 1957). A very large number of Staphylinidae, Catopidae, Diptera and Acarina were collected from the carcases of bank voles (Nabagło 1973). Watson in Louisiana, USA, collected in pitfall traps under six alligators, three bears, six deer and six swine a total of 218,514 Parasitidae mites (Watson 2004). During the fresh stage of decomposition 23 Parasitidae plus 7 seed mites, during the bloating stage 1,427 Parasitidae plus 99 seed, 7 needlenose, 4 mushroom and 2 strawberry mites, during active decomposition 5,062 Parasitidae plus 87 seed and 23 needlenose mites, during advanced decomposition 51,418 Parasitidae plus 104 seed and 6 needlenose mites and during dry decomposition 160,584 Parasitidae plus 194 seed, 15 needlenose, 8 strawberry and 4 mushroom mites. Unfortunately, the identity of the mites behind these vernacular names remains unresolved. For his twelfth case, Mégnin concluded: ‘the abundance of the Acarina, which were of an immense number, incalculable, on the leg of the mummy that we had to examine, proves that they were the principal agents of this mummification, without denying, however, that the abundance was helped by special environmental circumstances’ (Mégnin 1895). Von Niezabitowski (1902) also reported to always find larger numbers of mites belonging to the ‘Gamasidae’ (Mesostigmata) on human corpses but didn’t consider it to be characteristic. Mégnin’s first discovery of mites on and in a mummified newborn baby from the Paris area was followed by a report of a similar case from Montpellier in France (Brouardel 1879; Lichtenstein et al. 1885). The early cases describe the mummified corpses to be covered by a brownish layer some 2 mm thick and made up exclusively of mite carcases, exuvia and faeces (Brouardel 1879; Perotti and Braig 2009b). Such a brownish layer has been reported from many more cases of mummified corpses of babies and adults. However, in many cases this layer was not microscopically examined and the possible presence of mites was not detected (Strauch 1928; Forbes 1942). The detection of the small black fly Phora aterrima (Phoridae) in such a brownish layer might distract from looking for mites. When baby pig carcases were put in burial pits, during the later part of advanced decomposition, mites became so numerous that they gave the carcase a mottled appearance; and during dry decomposition, ants, flies, Collembola and mites were the dominant fauna (Payne et al. 1968). Myriads of mites, Thysanura (now order Collembola) and dipteran puparia but no beetles nor dipteran larvae were found on a human corpse interred for 4 years only in a burial case but without coffin in a grave 3 feet deep (Motter 1898). In a more recent case, the corpse of a young female recently exhumed after 28 years yielded thousands of live Collembola together with large numbers of Acari (mites) of the family Glycyphagidae, and fly puparia (Merritt et al. 2007). The only habitat where mites don’t seem to be numerous is on submerged carcases. In a study with baby pigs by Vance and colleagues, it was observed that during the collection 123 54 Exp Appl Acarol (2009) 49:45–84 process water mites and mayflies were typically found while searching the net holding the carcase after the net and carcase were recovered from submersion in a lake (Vance et al. 1995). The water mites detached readily during the first signs of carcase disturbance. In this study water mites were recovered in nine collections compared to amphipods in 19, mayflies in 20 and chironomids in 30 collections. However, Proctor expects freshwater mites to be of little forensic value in the estimation of post mortem intervals of submerged carcases (Proctor 2009). Buried carcases Corpses buried in graves only experience 4 waves of arthropod invasion (Mégnin 1887, 1894). In the introduction to the section on the fauna of buried and entombed corpses, Mégnin placed Acari next to Diptera, Coleoptera and Lepidoptera as constituents but did not elaborate further on any mite species that might be part of it, though he emphasised that the larvae of the mites were not visible to the naked eye. For the fourth and last wave of buried cases, the mite genera Uropoda and Trachynotus have been reported in the early literature (Lecha-Marzo 1917). A total of 150 exhumations in the late eighteenth century in Washington, DC (USA) yielded eight mite species on 30 human corpses, interred from 3 to 71 years (Motter 1898). This is a very high recovery rate for mites compared with insect taxa. The highest recovery rate was achieved for rove beetles of the genus Eleusis (Staphilinidae, Coleoptera), which were found in 56 cases interred from 1 to 11 years, followed by scuttle flies (Phoridae, Diptera), which were found on 43 human corpses interred from 3 to 38 years. The most commonly found mite species was the new species Uropoda depressa (Uropodidiae, Mesostigmata) present on bodies interred from 3 to 11 years. Again, this species new to science has not yet been systematically evaluated by acarologists. A completely dry and crumpling corpse interred for 71 years in a wood coffin 1.8 m deep in sandy soil contained no insects; only ‘Hypopus’ species, i.e. phoretic deutonynphs of several species in the family Acaridae (Astigmata) and a single snail, Helicodiscus lineatus, were present. In more recent exhuminations in France of shorter burial time, mites were reported from 3 of 22 human corpse, all in the stage of putrefaction and interred for 7–9 months (Bourel et al. 2004). Remarkably, conservation treatment applied to one of the corpses had no effect on the mite colonisation. Similarly, mites, springtails and puparia of coffin fly, Conicera tibialis, were collected from the embalmed body of a 28 year-old female with a gunshot wound to the head. The corpse was buried at a depth of 1.8 m in an unsealed casket that was placed inside an unsealed cement vault in a cemetery in Michigan, USA (Merritt et al. 2007). Mites in decomposition studies Mites have been observed in many decomposition studies but often referred to as Acari, Acarina or Acarida, for example: rabbits (Chapman and Sankey 1955), active and advanced decomposition, dry remains (Wolff et al. 2004); lizards and toads (Cornaby 1974); guinea pigs (Porta 1929); chickens, during all four or five stages of decomposition (Arnaldos et al. 2004; Horenstein et al. 2005); sparrows (Dahl 1896); pigs (Anderson et al. 2002; Grassberger and Frank 2004; Pérez et al. 2005; Schoenly et al. 2005; Kelly 2006); water mites on submerged pigs (Vance et al. 1995); sheep (Fuller 1934); mice and slugs (Kneidel 1984); voles (Nabagło 1973); crows, sparrows, striped field mice and baby pigs 123 Exp Appl Acarol (2009) 49:45–84 55 (Fourman 1936); a study involving some 1,200 rodent carcases in Wytham Woods around Oxford (Putman 1978); herring gulls and great black-backed gulls (Lord and Burger 1984b); fish (Walker 1957; Watson 2004); mites of the family Parasitidae on wild bear, deer, alligator and wild pig carcases (Watson and Carlton 2003). Mites have also been noticed at crime scenes or associated with human corpses but not identified (Bianchini 1929; Magni et al. 2008). In a study of the decomposition of baby pigs in Tennessee, USA, a total of 522 species representing 3 phyla, 9 classes, 31 orders, 151 families and 359 genera were identified (Payne 1965). Due to the need for a wide variety of taxonomic expertise, there is a tendency to report only a portion of the insects found on carrion based on the insect taxa previously published as forensically significant. This leads to a bias towards large, easily collected arthropods and avoidance of taxonomically difficult groups, i.e. Acari, Sphaeroceridae, Sepsidae, Histeridae, Drosophilidae, Piophilidae and many Staphylinidae (Gill 2005). This is also evident in the list of arthropod waves in Table 1, where authors indicated families instead of species. It is obvious that Acari—not being insects—should be the most difficult group of all for (forensic) entomologists. An extreme but fascinating case might demonstrate that even for arachnologists it might not be trivial to recognize a mite as such. Brucharachne ecitophila was initially described from a female specimen as the sole representative of the spider family Brucharachnidae. Reexamination revealed that the female spider specimen is actually a male dermanyssiod mite, now known as Sphaeroseius ecitophilus (Laelapidae, Mesostigmata) (Krantz and Platnick 1995). Along with size, the taxonomic difficulty of Acari might be the most important reason why mites are so often not reported in forensic and ecological studies of decomposition. Mites are part of a food web There are many ecological reasons why mites might be found on carcases. Mites will feed on successive waves of bacteria, algae and fungi that develop on the carcase. ‘Cheese’ mites that can be found feeding on cheese and ham, will feed on the caseous stage of carcases. Carcases pre-date cheese and ham in evolutionary terms. Species of macrochelid, parasitid, parholaspidid, uropodid and other mite families will prey on other mites, insects, and nematodes on the corpse. Nematodes have long been recognised as an integral part of animal and human decomposition but have been almost completely ignored by the forensic sciences. These nematodes, like the bacteria, algae and fungi, attract predatory mites to a carcase and then become as much part of the food web of a carcase as the nematodes. Other mite species specialise on the dry remains of the carcase. Several forensic web sources suggest that mites of the genus Rostrozetes (Haplozetidae, Oribatida) feed on dry skin in the later stages of decomposition. While a large diversity of mite species has been collected at later stages of decomposition and from dry skin (Table 3), there is currently no evidence for any Rostrozetes species being associated with animal or human remains. Several species of Rostrozetes are very common inhabitants of leaf litter and peatlands and are found on moss and fungi from tree trunks (Behan-Pelletier and Bissett 1994). Reports on associations of Rostrozetes with animal skin are very rare and restricted to parasitic infestations of living animals (Parker and Holliman 1971). Burying and sexton beetles (Nicrophorus spp., Silphidae) bring mites of the genus Poecilochirus (Parasitidae, Mesostigmata) to a carcase. These mites have long been implicated in a symbiotic interaction with their carrier host. Poecilochirus can kill the eggs of blow flies, which are one of the main competitors of these beetles for the carcase. Blow 123 Species Family Abundance Host Location Habitat Country Season (month) Reference On On Under On, under Grassland, bush Grassland, bush Woods Xero ? mesophytic Spain Spain ME, USA HI, USA 12–2 12–2 Castillo Miralbes 2002 Castillo Miralbes 2002 Wasti 1972 Early and Goff 1986; Goff 1989 56 123 Table 3 Within the decompositional stages, families with mite species reported from human corpses are listed first followed in alphabetical order by other families reported only from animal carcases Fresh stage, initial decay Mesostigmata Arctoseius sp. Haemogamasus sp. Glyptholaspis americana Macrocheles merdarius M. muscaedomesticae Parasitus sp. Poecilochirus necrophori P. silphaphila Increasing Some Pig Pig Chicken Cat Macrochelidae Some Some Some Cat Cat Pig On, under On, under On, under Xero ? mesophytic HI, USA Xero ? mesophytic HI, USA Several HI, USA Increasing High Chicken Pig Mice Under On On Woods Grassland, bush Forest Increasing Abundant Some Some Large carcases Chicken Chicken Cat Pig On Under On On, under On MI, USA Woods ME, USA Farm IA, USA Xero ? mesophytic HI, USA Several HI, USA Dog Assoc. Parasitidae Parasitidae Urodinychidae Uropodidae ME, USA Spain MI, USA 3–5 3–5 3–5 12–2 3–5 Early and Goff 1986; Goff 1989 Early and Goff 1986; Goff 1989 Richards and Goff 1997; Avila and Goff 1998; Davis and Goff 2000 Wasti 1972 Castillo Miralbes 2002 Wilson 1983 Yoder 1972; Brown and Wilson 1994 Wasti 1972 Rives and Barnes 1988 Early and Goff 1986; Goff 1989 Hewadikaram and Goff 1991; Avila and Goff 1998 Astigmata Acarus farris Acaridae A. siro Acaridae Acaridae Common Moderate Lizard, chicken On Chicken Under Chicken On, under Woods Woods Field Costa Rica NY, USA Nigeria MA, USA Spain 10–3 OConnor 2009 OConnor 2009 Iloba and Fawole 2006 Wasti 1972 Arnaldos et al. 2004 Exp Appl Acarol (2009) 49:45–84 Fuscuropoda sp. Sp 1–3 Ascidae Haemogamasidaea Laelapidae Macrochelidae Species Family Abundance Host Location Habitat Country Season (month) Reference Oribatids Decline Chicken Under Woods MA, USA Wasti 1972 Demodecidae Decline Decline Decline Abundant Abundant Moderate Human Human Most mammals Chicken Chicken Chicken Pig On On On Under Under Under On Normal fauna Normal fauna Normal fauna Woods Woods Woods Bush Worldwide Worldwide Worldwide MA, USA MA, USA MA, USA Spain Desch 2009 Desch 2009 Wilson 1844; Gmeiner 1908 Wasti 1972 Wasti 1972 Wasti 1972 Castillo Miralbes 2002 On On Under On, under Grassland, bush Grassland, bush Woods Xero ? mesophytic Spain Spain ME, USA HI, USA Oribatida Prostigmata Demodex brevis D. folliculorum Demodecidae Bdellidae Rhagidiidae Trombidiidae 12–2 Exp Appl Acarol (2009) 49:45–84 Table 3 continued Putrefaction, bloated stage—terrestrial Mesostigmata Arctoseius sp. Haemogamasus sp. Glyptholaspis americana Macrocheles merdarius M. muscaedomesticae Pachylaelaps sp. Parasitus sp. Pergamasus sp. Fewer Some Pig Pig Chicken Cat Macrochelidae Some Some Some Cat Cat Pig On, under On, under On, under Xero ? mesophytic HI, USA Xero ? mesophytic HI, USA Several HI, USA Fewer Some Chicken Cat Pig Cat Pig Chicken Pig Under On, under On On, under On, under Under On, under Woods Xero ? mesophytic Grassland, bush Xero ? mesophytic Several Woods Several Macrochelidae Pachylaelapidae Parasitidae Parasitidae Some Some Fewer Common ME, USA HI, USA Spain HI, USA HI, USA ME, USA HI, USA 1–3 1–3 3–5 3–5 3–5 3–5 1–3 3–5 Castillo Miralbes 2002 Castillo Miralbes 2002 Wasti 1972 Early and Goff 1986; Goff 1989 Early and Goff 1986; Goff 1989 Early and Goff 1986; Goff 1989 Richards and Goff 1997; Avila and Goff 1998; Davis and Goff 2000 Wasti 1972 Early and Goff 1986; Goff 1989 Castillo Miralbes 2002 Early and Goff 1986; Goff 1989 Hewadikaram and Goff 1991 Wasti 1972 Richards and Goff 1997; Avila and Goff 1998 57 123 Ascidae Haemogamasidae Laelapidae Macrochelidae 58 123 Table 3 continued Species Family Abundance Host Location Habitat Country Season (month) Reference Poecilochirus sp. Sp 1–3 Uropodidae Some Some Some Rabbit Cat Pig On On, under On Woods CO, USA Xero ? mesophytic HI, USA Several HI, USA 7–8 3–5 De Jong and Chadwick 1999 Early and Goff 1986; Goff 1989 Hewadikaram and Goff 1991; Avila and Goff 1998 Acarus siro Acaridae Common Woods Lardoglyphus zacheri Lardoglyphidae or Acaridae Fish, frog, Lizard, chicken On Deer Assoc. Nigeria UT, USA Pig On Grassland, bush Spain 9,1–3 Castillo Miralbes 2002 Pig On Bush Spain 5 Castillo Miralbes 2002 Pig On One Human (26 d) On Forest Ascidae Haemogamasidae Macrochelidae Abundant Pig Pig Cat On On On, under Macrochelidae Abundant Abundant Cat Pig Pachylaelapidae 20, 2 Abundant Rat Cat Astigmata Iloba and Fawole 2006 OConnor 2009 Prostigmata Trombidiidae Ixodida Ixodidae Putrefaction, bloated stage—freshwater Oribatida Hydrozetes sp. Hydrozetidae Black putrefaction, active decay Hobischak and Anderson 2002 Canada Mesostigmata Macrocheles sp. Pachylaelaps sp. Rhodacaridae 12 Leclercq 1978 Grassland, bush Spain Grassland, bush Spain Xero ? mesophytic HI, USA 2–4 2–4 3–5 Castillo Miralbes 2002 Castillo Miralbes 2002 Early and Goff 1986; Goff 1989 On, under On, under Xero ? mesophytic HI, USA Several HI, USA 3–5 On, under On, under Germany Xero ? mesophytic HI, USA Early and Goff 1986; Goff 1989 Richards and Goff 1997; Avila and Goff 1998; Davis and Goff 2000 Schönborn 1963 Early and Goff 1986; Goff 1989 Belgium 3–5 Exp Appl Acarol (2009) 49:45–84 Cyrtolaelaps mucronatus Arctoseius sp. Haemogamasus sp. Glyptholaspis americana Macrocheles merdarius Species Family Parasitus stercorarius Parasitidae Common Taxonomic position uncertain Parasitus sp. Pergamasus sp.b Location Habitat Country Pig, sparrow, Crow, mouse On Forest Germany Pig Cat Pig Harbour seal Rabbit Pig On On, under On, under On On On, under Grassland, bush Xero ? mesophytic Several Rock Woods Several Spain HI, USA HI, USA MA, USA CO, USA HI, USA Season (month) Reference Fourman 1936 Castillo Miralbes 2002 Early and Goff 1986; Goff 1989 Hewadikaram and Goff 1991 5–10 Lord and Burger 1984a 7–8 De Jong and Chadwick 1999 Richards and Goff 1997; Avila and Goff 1998 Woods W Australia 10–12 Bornemissza, 1957 Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989 On Several HI, USA Hewadikaram and Goff 1991; Avila and Goff 1998 Parasitidae Abundant Abundant Common Common Abundant ‘Gamasidae’ Uropodidae Large Abundant Guinea pig Cat Abundant Under On, under Pig Acaridae Common Abundant Abundant Fish, frog, pig Cat Cat On, under On, under Xero ? mesophytic HI, USA Xero ? mesophytic HI, USA Abundant Abundant Pig Pig Common crow, White-tailed deer Mouse, crow, White–tailed deer Guinea pig On On Several Several Poecilochirus sp. Sp 1–3 Abundance Host 2–4 3–5 Exp Appl Acarol (2009) 49:45–84 Table 3 continued Astigmata Acarus siro Sancassania berlesi Tyrophagus putrescentiae Spinanoetus spp. nov Acaridae Histiostomatidae ‘Tyroglyphidae’ Some Early and Goff 1986; Goff 1989 Early and Goff 1986; Goff 1989 HI, USA HI, USA Hewadikaram and Goff 1991 Avila and Goff 1998 On MI, USA OConnor 2009 On MI, USA OConnor 2009 W Australia 10–12 Bornemissza 1957 Under Woods 59 123 Pelzneria spp. nov 3–5 3–5 60 123 Table 3 continued Species Family Abundance Host Location Habitat Country Season (month) Reference Arnaldos et al. 2004 Arnaldos et al. 2004 Oribatida Galumna tarsipennata Zygoribatula connexa Galumnidae Oribatulidae Chicken Chicken On, under On, under Field Field Spain Spain 2–3 2–3 Trombidiidae Pig On Grassland, bush Spain 9–10,3–4 Castillo Miralbes 2002 Pig On Grassland, bush Spain 2–5 Castillo Miralbes 2002 Scarce Many Pig Dog Human (3 m) On In, on, under On, under Grassland, bush Woods, pasture Deciduous forest Spain TN, USA Spain 10,4 1–12 8 Castillo Miralbes 2002 Reed 1958 Saloña et al. in prep. Ten Many Many Ten Many Human Human Human Human Human m) m) m) m) m) On Under On, under On Under Belgium Spain Spain Belgium Spain 12 8 8 12 8 Leclercq and Verstraeten 1988b Saloña et al. in prep. Saloña et al. in prep. Leclercq and Verstraeten 1988b Saloña et al. in prep. Common Cat Fox Cat Human (17 d) Impala On, under On On, under On On 3–5 10 3–5 10 1–10 Early and Goff 1986; Goff 1989 Smith 1975 Early and Goff 1986; Goff 1989 Easton and Smith 1970 Braack 1986, 1987 Cat Dog Chicken Pig On, under In, on, under Under On 3–5 1–12 Early and Goff 1986; Goff 1989 Reed 1958 Wasti 1972 Payne and Crossley 1966 Prostigmata Ixodida Ixodidae Butyric fermentation, advanced decay Mesostigmata Macrocheles glaber M. merdarius M. muscaedomesticae Macrocheles sp. Ascidae Laelapidae Macrochelidae Common One Some Common Abundant Increase (2 (3 (3 (2 (3 Deciduous forest Deciduous forest Deciduous forest Xero ? mesophytic Garden Xero ? mesophytic Small wood Woods HI, USA England HI, USA England South Africa Xero ? mesophytic HI, USA Woods, pasture TN, USA Woods ME, USA Woods SC, USA 8 Exp Appl Acarol (2009) 49:45–84 Arctoseius sp. Asca sp. Proctolaelaps epuraeae Proctolaelaps sp. ? Zerconopsis remiger Hypoaspis aculeifer Hypoaspis sp. ? Glyptholaspis americana Species Gamasodes spiniger ‘Gamasus’ sp. Paragamasus sp. Parasitus fimetorum Parasitus sp. Family Abundance Host Location Habitat Country Macrochelidae Common Pig On, under Several HI, USA Fox Human (2.7 y) Human (3 m) Fox Dog Chicken Pig Pig Dog Rabbit On On Under On In, on, under Under On On In, on, under On, under Garden Cask Deciduous forest Garden Woods, pasture Woods Grassland Woods Woods, pasture Urban On On beetle Under Pine forest Medium Human (35 d) Human (2 m) Human hanging Human (17 d) Human (35 d) Harbour seal Rabbit Guinea pig Human (3 m) Pig Dog England Switzerland Spain England TN, USA ME, USA Spain SC, USA TN, USA Alex., Egypt Belgium Spain On On On On Under On, under On In, on, under Rare Pig Dog On In, on, under Parasitidae Many Many Abundant Increase Pergamasus sp. Phorytocarpais spp. Scarce Abundant Poecilochirus carabi Common Several Many P. necrophori P. subterraneus Poecilochirus sp. Urobovella pulchella Uroseius sp. Apionoseius sp. Discourellidae Idendity unclear Haemogamasidae 10 8 10 1–12 4 8 1–12 11–4 8 11 Reference Richards and Goff 1997; Avila and Goff 1998; Davis and Goff 2000 Smith 1975 Hunziker 1919 Saloña et al. in prep. Smith 1975 Reed 1958 Wasti 1972 Castillo Miralbes 2002 Payne and Crossley 1966 Reed 1958 Tantawi et al. 1996 Leclercq and Verstraeten 1988b Saloña-Bordas pers. comm. Small wood Rock Woods Woods Deciduous forest Grassland, bush Woods, pasture England Belgium MA, USA CO, USA W Australia Spain Spain TN, USA 10 8 5–10 7–8 10–12 8 10 1–12 Easton and Smith 1970 Leclercq and Verstraeten 1988b Lord and Burger 1984a De Jong and Chadwick 1999 Bornemissza, 1957 Saloña et al. in prep. Castillo Miralbes 2002 Reed 1958 Grassland, bush Woods, pasture Spain TN, USA 10,4 1–12 Castillo Miralbes 2002 Reed 1958 61 123 Haemogamasus sp. Melittiphis ? sp. ‘Gamasidae’ Uropodidae One Common Common Common Large Many Season (month) Exp Appl Acarol (2009) 49:45–84 Table 3 continued 62 123 Table 3 continued Species Family Abundance Host Location Habitat Country Season (month) Reference Gamasellus sp. Rhodacaridae or Ologamasidae Zerconidae Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958 Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958 Myianoetus diadematus Histiostomatidae Acarus siro Acaridae Mass occ. Common On Basement Germany Cosmoglyphus sp. Sancassania berlesei Common Many Common Common Human (1.3 y) Fish, frog, pig Lizard, chicken Cat Human (3 m) Cat Cat Deer, raccoon Dog Pig Cat On On, under Under On, under On, under On Under On On, under Woods Xero ? mesophytic Deciduous forest Xero ? mesophytic Xero ? mesophytic Several HI, USA TX, USA Hewadikaram and Goff 1991 OConnor 2009 Woods Several W Australia 10–12 HI, USA Bornemissza 1957 Avila and Goff 1998 Zercon sp. Astigmata Sancassania sp. nov Sancassania sp. nov Sancassania sp. Few Abundant Common Tyrophagus putrescentiae Lardoglyphus zacheri 3–5 8 3–5 3–5 1–12 6–11 3–5 Iloba and Fawole 2006 Early and Goff 1986; Goff Saloña et al. in prep. Early and Goff 1986; Goff Early and Goff 1986; Goff OConnor 2009 Reed 1958 Payne et al. 1968 Early and Goff 1986; Goff 1989 1989 1989 1989 Common Pig Bird Immense Common Guinea pig Pig On Feathers under Under, on On Many Few Many Human (3 m) Dog Human (3 m) Under Under Under Deciduous forest Woods, pasture Deciduous forest Spain TN, USA Spain 8 1–12 8 Saloña et al. in prep. Reed 1958 Saloña et al. in prep. Few Few Common Dog Dog Pig Chicken Under Under On On, under Woods, pasture Woods, pasture Several Field TN, USA TN, USA HI, USA Spain 1–12 1–12 Reed 1958 Reed 1958 Hewadikaram and Goff 1991 Arnaldos et al. 2004 Oribatida Platynothrus peltifer Camisiidae Minunthozetes semirufus Malacanothrus sp. Ceratoppia bipilis Mycobatidae Medioppia pinsapi Malacanothridae Ceratoppiidae Liacaridae Oppiidae 10–12 Exp Appl Acarol (2009) 49:45–84 Lardoglyphidae or Acaridae ‘Tyroglyphidae’ Acaridae Nigeria HI, USA Spain HI, USA HI, USA USA Woods, pasture TN, USA Burial pit SC, USA Xero ? mesophytic HI, USA Russell et al. 2004 Species Family Oribatula tibialis Oribatulidae Oribatida spp. Abundance Host Location Habitat Country Season (month) Reference Chicken Pig Dog Dog On, under On Under Under Field Several Woods, pasture Woods, pasture Spain HI, USA TN, USA TN, USA 10–12 1–12 1–12 Arnaldos et al. 2004 Davis and Goff 2000 Reed 1958 Reed 1958 Pig Dog Pig On Under On Woods Woods, pasture Grassland, bush SC, USA TN, USA Spain 8 1–12 10 Payne and Crossley 1966 Reed 1958 Castillo Miralbes 2002 Few Dog Pig Under On Woods, pasture Bush TN, USA Spain 1–12 4 Reed 1958 Castillo Miralbes 2002 Myriads Human ([1 y) On, in Cellar France Mégnin 1894 Common Common Common On On On Grave Grave Canada DC, USA DC, USA Johnston and Villeneuve 1897 Motter 1898 Motter 1898 Common Common Rare Common Human Human (11 y) Human (20–30 y) Dog (3 m) Human (11 y) Dog Human (53 d) On On In, on, under Soil Grave Grave Woods, pasture Clothing DC, USA DC, USA TN, USA HI, USA 1–12 3–5 Motter 1898 Motter 1898 Reed 1958 Goff 1991 Common One Common Common Cat Human (3 m) Human (53 d) Cat On, under On Soil On, under Xero ? mesophytic HI, USA Belgium Clothing HI, USA Xero ? mesophytic HI, USA 6 3–5 3–5 Early and Goff 1986; Goff 1989 Leclercq and Verstraeten 1988b Goff 1991 Early and Goff 1986; Goff 1989 Common Few Few Prostigmata Lephus spp. Penthaleus major Erythraeidae Eupodidae Trombidiidae Few Exp Appl Acarol (2009) 49:45–84 Table 3 continued Ixodida Dermacentor variabilis Ixodidae Ixodidae Dry decomposition, skeletal stage Mesostigmata Leiodinychus krameri Dinychidae (Uropodidae) Holostaspis sp. Hypoaspis sp. Laelapidae Laelaps (Iphis) sp. Melittiphis ? sp. Glyptholaspis americana 63 123 Macrocheles glaber M. merdarius Macrochelidae 64 123 Table 3 continued Species Family Abundance Host Location Habitat Country Season (month) Reference Cat Impala Dog Small animalsf Pig On, under On In, on, under On On, under Xero ? mesophytic Woods Woods, pasture Oak forest Several HI, USA Sth Africa TN, USA IL, USA HI, USA 3–5 1–10 1–12 4–11 Macrochelidae Common Some Abundant Abundant Common Common Common Common Human (53 d) Cat Human (30–40 y) Dog (3 m) Dog Small animalsf Dog Small animalsf Rabbit Human (3 y) Human (3–7 y) Soil On, under On Clothing HI, USA Xero ? mesophytic HI, USA Grave DC, USA Early and Goff 1986; Goff 1989 Braack 1986, 1987 Reed 1958 Johnson 1975 Richards and Goff 1997; Avila and Goff 1998; Davis and Goff 2000 Goff 1991 Early and Goff 1986; Goff 1989 Motter 1898 On In, on, under On In, on, under On On On, in On Grave Woods, pasture Oak forest Woods, pasture Oak forest Woods Grave DC, USA TN, USA IL, USA TN, USA IL, USA CO, USA France DC, USA Common Common Common Dog (3–5 m) Human (53 d) Cat Common On Soil On, under Pig Grave Clothing Xero ? mesophytic On DC, USA HI, USA HI, USA Several Scarce Medium Dog Dog In, on, under In, on, under Woods, pasture Woods, pasture TN, USA TN, USA Motter 1898 3–5 Goff 1991 3–5 Early and Goff 1986; Goff 1989 HI, USA Hewadikaram and Goff 1991; Avila and Goff 1998 1–12 Reed 1958 1–12 Reed 1958 Rare Dog In, on, under Woods, pasture TN, USA 1–12 M. muscaedomesticae Macrocheles sp. Pachylaelaps sp. Pachylaelapidae ‘Gamasus’ sp. Parasitidae Parasitus sp. Pergamasus sp.c Poecilochirus sp. Uroseius acuminatus Uropoda depressa Asca sp. Apionoseius sp. Gamasellus sp. Uropodidae Ascaidae Discourellidae Identity unclear Rhodacaridae or Ologamasidae 1–12 4–11 1–12 4–11 7–8 5 Motter 1898 Reed 1958 Johnson 1975 Reed 1958 Johnson 1975 De Jong and Chadwick 1999 Mégnin 1894 Motter 1898 Reed 1958 Exp Appl Acarol (2009) 49:45–84 Uropoda sp. Sp 1–3 Trachytidae Uropodidae Identity unclear Common Abundant Common Scarce Abundant Common Common Common 3–5 3–5 Species Family Abundance Host Location Habitat Country Season (month) Reference Zercon sp. Zerconidae Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958 Acaridae Common On On On, in On, in Basement Rural Cellar Germany OH, USA France France On On Woods Grave Nigeria DC, USA Astigmata A. siro Common Myriads Common Acarus (Tyroglyphus) sp. Cosmoglyphus sp. Rhizoglyphus echinopus Common Human (1.3 y) Raccoon Human (3 y) Human ([1 y) Fish, frog, pig Lizard, chicken Human (3 y) Common Myriads Cat Human ([1 y) On, under On, in Xero ? mesophytic HI, USA Cellar France 3–5 Early and Goff 1986; Goff 1989 Mégnin 1894 Abundant Common Abundant Human (2 y) Human (2–3 y) Human (3–8 m) Human (7–8 y) Human (3 m) Human (3.5 m) Cat Cat Deer, raccoon Dog Human (3–8 m) Human (7–8 y) Human (1 y) On Bulbs of lily On, in Urban Garden, burrial Urban 10 Mégnin 1894 Mégnin 1894 Brouardel 1879 On On On On, under On, under On Under On, in House France Belgium Belgium Xero ? mesophytic HI, USA Xero ? mesophytic HI, USA USA Woods, pasture TN, USA Urban France On On House House Acarus immobilis Sancassania berlesei Sancassania sp. nov Sancassania sp. nov Sancassania sp. Tyrophagus longior Abundant 784 Abundant Common Common Few Abundant France France Iloba and Fawole 2006 Motter 1898 1 1–12 1 Mégnin 1894 Leclercq and Verstraeten 1988b Leclercq and Verstraeten 1988b Early and Goff 1986; Goff 1989 Early and Goff 1986; Goff 1989 OConnor 2009 Reed 1958 Brouardel 1879 1 Mégnin 1894 Mégnin 1894 6 1 3–5 3–5 65 123 Abundant Very rare France France France 5 Russell et al. 2004 OConnor 2009 Mégnin 1894 Mégnin 1894 Exp Appl Acarol (2009) 49:45–84 Table 3 continued 66 123 Table 3 continued Species Family T. putrescentiae Tyrophagus sp. T. (Hypopus) sp. Abundance Host Location Habitat Country Common Myriads Myriads Common Abundant Common Common Common Common Common Common Human (3 y) Human ([1 y) Human Human Human (1.3 y) Human (53 d) Cat Pig Human (18 m) Human (2 y) Human (20–71 y) On, in On, in On On, in On Soil On, under On On, around On On Rural Cellar House, trunk Acaridae Few Alps France Common Very rare Common Large Common Common Common Common Pig Human (1 y) Human Human (28 y) Human Human Human Human Human (summer) On On On On On On On On Several House HI, USA France France MI, USA France Canada France Canada Two One Common Common Human (3 m) Human (3 m) Human (53 d) Cat On On Soil On, under Belgium Belgium Clothing HI, USA Xero ? mesophytic HI, USA France France France Canada Basement Germany Clothing HI, USA Xero ? mesophytic HI, USA Several HI, USA House France Urban France Grave DC, USA Season (month) Reference 5 Mégnin 1894 Mégnin 1894 Mégnin 1898 Johnston and Villeneuve 1897 Russell et al. 2004 Goff 1991 Early and Goff 1986; Goff 1989 Hewadikaram and Goff 1991 Mégnin 1894 Mégnin 1894 Motter 1898 3–5 3–5 3 6 Identity unclear Glycyphagus destructor Glycyphagidae H. necrophagus Composite species, unrecognisable H. sachsi Histiostoma sp. Embalmed 1 6 6 3–5 3–5 Leclercq and Verstraeten 1992 Avila and Goff 1998 Mégnin 1894 Mégnin 1894 Merritt et al. 2007 Mégnin 1894 Johnston and Villeneuve 1897 Mégnin 1894 Johnston and Villeneuve 1897 Leclercq and Verstraeten 1988b Leclercq and Verstraeten 1988b Goff 1991 Early and Goff 1986; Goff 1989 Exp Appl Acarol (2009) 49:45–84 Glycyphagidae Histiostoma feroniarum Histostomatidae 10 Species Family Abundance Host Location Habitat Histostomatidae Some Common Cat Pig On, under On, under Xero ? mesophytic HI, USA Several HI, USA Human Pelvis, Mummy Common Human Cat Racoon Human (53 d) Gut content Gut content On, under On Soil Mummy Chile Xero ? mesophytic HI, USA USA Clothing HI, USA Common Cat On, under Xero ? mesophytic HI, USA Common Few Few Some Common 238 Common Few Common Few Increase Human Human Dog Dog Cat Skin of animals Rat Pig Dog Pig Dog Chicken Remains On Under Under On, under On On On Under On Under Under Tomb Grave Woods, pasture Woods, pasture Xero ? mesophytic Spain DC, USA TN, USA TN, USA HI, USA Campus Several Woods, pasture Several Woods, pasture Woods Cameroon HI, USA TN, USA HI, USA TN, USA MA, USA Abundant Human ([1 y) On Cellar France Myianoetus ? sp. Lardoglyphus radovskyi Lardoglyphidae or Acaridae L. robustisetosus L. zacheri Czenspinskia transversostriata Some Winterschmidtiidae Country Season (month) Reference 3–5 Early and Goff 1986; Goff 1989 Richards and Goff 1997; Avila and Goff 1998 Baker 1990 NV, USA 3–5 Radovsky 1970 Baker 1990 Early and Goff 1986; Goff 1989 OConnor 2009 Goff 1991 3–5 Early and Goff 1986; Goff 1989 1–12 1–12 3–5 Hidalgo-Argüello et al. 2003 Motter 1898 Reed 1958 Reed 1958 Early and Goff 1986; Goff 1989 3–5 Exp Appl Acarol (2009) 49:45–84 Table 3 continued Oribatida Aphelacarus acarinus Hoplophora (Tritia) sp. Platynothrus peltifer Ceratoppia bipilis Rostrozetes spp.d Malacanothrus sp. Aphelacaridae Euphthiracaridae Camisiidae Ceratoppiidae Galumnidae Haplozetidae Haplozetidae Liacaridae Malacanothridae Oribatida spp. 2–3 1–12 1–12 Feugang Youmessi et al. 2008 Hewadikaram and Goff 1991 Reed 1958 Davis and Goff 2000 Reed 1958 Wasti 1972 Cheyletus eruditus Cheyletidae Mégnin 1894 67 123 Prostigmata 68 123 Table 3 continued Species Family Tarsotomus sp. nov Podapolipidae Tarsonemoidea Anystidae Erythraeus sp. Penthaleus major Trombidium sp. Erythraeidae Eupodidae Trombidiidae Abundance Host Location Habitat Country Abundant Human Human Rabbit Remains Remains On, under Tomb Tomb Urban Few Common Pig Dog Small animalsf On Under On Woods Woods, pasture Oak forest Spain Spain Alexandria, Egypt SC, USA TN, USA IL, USA Few Dog Pig Under On Woods, pasture Woods Male Small animal General, human? General, human? Small animal Small animal On On Alder forest Season (month) Reference 7–8 Hidalgo-Argüello et al. 2003 Hidalgo-Argüello et al. 2003 Tantawi et al. 1996 8 1–12 4–11 Payne and Crossley 1966 Reed 1958 Johnson 1975 TN, USA SC, USA 1–12 8 Reed 1958 Payne and Crossley 1966 Poland 8 Gwiazdowicz and Klemt 2004 Porta 1929 Ixodida Dermacentor variabilis Ixodidae Undetermined stage Mesostigmata Epicrius mollis E. thanathophilus Epicriidae Celaenopsis cuspidatus Celaenopsidae Parasitidae Pergamasus crassipes Pergamasus sp.e Poecilochirus sp. Parasitidae Few Few Few Hundreds Decrease Common Common General, human? Small animal Rat Rat Rat Bear, deer, Alligator, pig On On Porta 1929 Alder forest Alder forest Poland Poland 8 8 On On On Under On On Gwiazdowicz and Klemt 2004 Gwiazdowicz and Klemt 2004 Porta 1929 Alder forest Copse, grassland Copse, grassland Field Poland England England CO, USA LA, USA 8 8–12 8–12 7–8 Gwiazdowicz and Klemt 2004 Collins 1970 Collins 1970 De Jong and Hoback 2006 Watson 2004 Exp Appl Acarol (2009) 49:45–84 Cornigamasus lunaris Holoparasitus calcaratus Paracarpais furcatus On Species Asca craneta Gamasellodes bicolor Iphidozercon gibbus Proctolaelaps sp. nov Zerconopsis decemremiger Hypoaspis (Cosmolaelaps) vacua Family Abundance Host Location Habitat Country Season (month) Reference Gamasida Ascidae Some Some Few Some Some Some Pig Cat Small animal Small animal Cat Small animal On On, under On On On, under On Several Xero ? mesophytic Alder forest Alder forest Xero ? mesophytic Alder forest HI, USA HI, USA Poland Poland HI, USA Poland 1–4 3–5 8 8 3–5 8 Davis and Goff 2000 Early and Goff 1986; Goff 1989 Gwiazdowicz and Klemt 2004 Gwiazdowicz and Klemt 2004 Early and Goff 1986; Goff 1989 Gwiazdowicz and Klemt 2004 Ascidae Some Pig On, under Several HI, USA Digamasellidae Digamasellidae Diplogyniidae Eviphidae Laelapidae Some As control Increasing Some Some Pig Turtle Turtle Pig Small animal On Under Under On On Several Woods Woods Several Alder forest HI, USA MA, USA MA, USA HI, USA Poland Laelapidae 146, large Increase Some Hundreds Turtle Rat Pig Rat Rodents Carrion Roe deer Weasel Fish, squid, carrion Vole Small mammals On, under Under Associated On On In On On In Woods Copse, grassland Several Copse, grassland MA, USA England HI, USA England USA Australia Slovakia MT, USA Australia Macrocheles sp. Macrochelidae Macrocheles sp. N M. agilis M. lagodekhensis group M. matrius M. mykytowyczi On On Forest Lithuania USSR 8 6–8 8–12 8–12 4, 5,7 Richards and Goff 1997; Avila and Goff 1998 Avila and Goff 1998 Abell et al. 1982 Abell et al. 1982 Avila and Goff 1998 Gwiazdowicz and Klemt 2004 Abell et al. 1982 Collins 1970 Richards and Goff 1997 Collins 1970 Krantz and Whitaker 1988 Halliday 2000 Mašán 1993 Krantz and Whitaker 1988 Halliday 2000 Hyatt and Emberson 1988 Bregetova and Koroleva 1960 69 123 M. nataliae (=melisii) Several 6–8 6–8 Exp Appl Acarol (2009) 49:45–84 Table 3 continued 70 123 Table 3 continued Species Family Abundance Host Location Habitat Country Carrion trap Squid Squid Rat Pig Pig Cat Turtle Pig Rat Small animal Turtle Rat Small animal Pig On On On Under Associated Associated On, under Far, under Associated Under On Under Under On On Australia Australia Australia Copse, grassland England Several HI, USA Several HI, USA Xero ? mesophytic HI, USA Woods MA, USA Several HI, USA Copse, grassland England Alder forest Poland Woods MA, USA Copse, grassland England Alder forest Poland Several HI, USA Season (month) Reference Veigaia nemorensis Veigaiidae Veigaiidae Increase Some Some Some Increasing Some Decrease Several As control Increase Few Some Prozercon kochi Zerconidae Few Small animal On Alder forest Poland 8 Halliday 2000 Halliday 2000 Halliday 2000 Collins 1970 Richards and Goff 1997 Richards and Goff 1997 Early and Goff 1986; Goff 1989 Abell et al. 1982 Richards and Goff 1997 Collins 1970 Gwiazdowicz and Klemt 2004 Abell et al. 1982 Collins 1970 Gwiazdowicz and Klemt 2004 Richards and Goff 1997; Davis and Goff 2000 Gwiazdowicz and Klemt 2004 Acaridae Lardoglyphidae or Acaridae Astigmata As control Some Turtle Impala Under On Woods Woods 6–8 1–10 Abell et al. 1982 Braack 1986 Decrease Rat Under Copse, grassland MA, USA South Africa England 8–12 Collins 1970 Achipteriidae Ceratozetidae Decrease Increasing Decrease One Decrease Rat Turtle Rat Rat Rat Under Under Under On Under Copse, grassland Woods Copse, grassland Field Copse, grassland England MA, USA England CO, USA England 8–12 6–8 8–12 7–8 8–12 Collins 1970 Abell et al. 1982 Collins 1970 De Jong and Hoback 2006 Collins 1970 M. peckorum M. spatei M. tessellatus Macrochelidae Ologamasidae Paraholaspidae Phytoseius hawaiiensis Phytoseiidae Phytoseiidae Podocinidae Rhodacaridae Trachytes aegrota Trachytidae Uropodidae 8–12 3–5 6–8 8–12 8 6–8 8–12 8 Astigmata Oribatida Eremaeidae Euzetidae Exp Appl Acarol (2009) 49:45–84 Lardoglyphus sp. Species Family Abundance Host Location Habitat Country Season (month) Reference Galumnidae Hypochthoniidae Mycobatidae Nothridae Increasing Increasing Decrease 3, small Decrease Decrease Decrease Turtle Turtle Rat Turtle Rat Rat Rat Far, under Far, under Under On, under Under Under Under Woods Woods Copse, grassland Woods Copse, grassland Copse, grassland Copse, grassland MA, USA MA, USA England MA, USA England England England 6–8 6–8 8–12 6–8 8–12 8–12 8–12 Abell et al. 1982 Abell et al. 1982 Collins 1970 Abell et al. 1982 Collins 1970 Collins 1970 Collins 1970 On Few Some Some Some Some Some Some Some General, human? Rat Pig Pig Cat Cat Pig Pig Impala Scutacaridae Tarsonemoidea Prostigmata Some Some Increasing Some Some Some Decrease Cat Pig Turtle Pig Cat Pig Rat On, under On Under Associated On, under On Under Field CO, USA Several HI, USA Several HI, USA Xero ? mesophytic HI, USA Xero ? mesophytic HI, USA Several HI, USA Several HI, USA Woods South Africa Xero ? mesophytic HI, USA Several HI, USA Woods MA, USA Several HI, USA Xero ? mesophytic HI, USA Several HI, USA Copse, grassland England Terpnacaridae Some Pig On Several Oppiidae Oribatulidae Exp Appl Acarol (2009) 49:45–84 Table 3 continued Prostigmata Erythraeus sabulosus Cunaxa sp. Pygmephorus sp. Erythraeidae Anystidae Bdellidae Camerobiidae Cunaxidae Ereynetidae Eupodidae Pygmephoridae Pygmephoridae Rhagidiidae On Associated On On, under On, under On Associated On Porta 1929 7–8 3–5 3–5 1–10 3–5 6–8 3–5 Early and Goff 1986; Goff 1989 Avila and Goff 1998 Abell et al. 1982 Richards and Goff 1997 Early and Goff 1986; Goff 1989 Avila and Goff 1998 Collins 1970 Endeostigmata HI, USA Avila and Goff 1998 71 123 8–12 De Jong and Hoback 2006 Richards and Goff 1997 Avila and Goff 1998 Early and Goff 1986; Goff 1989 Early and Goff 1986; Goff 1989 Avila and Goff 1998 Richards and Goff 1997 Braack 1986 72 123 Table 3 continued Species Family Abundance Host Location Few On Habitat Country Season (month) Reference LA, USA 12–2 Watson 2004 Ixodida Dermacentor variabilis Ixodidae Bear In many reports, particular mites species or taxa dominate in a single stage of decomposition but are also recorded in lower numbers in the adjacent preceding and subsequent stage. To increase clarity, the presence in these adjacent stages is not recorded in the table unless the species is dominant in more than one stage. The description of abundance tries to follow the original wording of the authors. If mites are abundant, one might expect several hundred thousand to several million specimens per carcase. For a few cases of human corpses, the estimated post mortem interval is given between parentheses in the column of host designation and the month the corpse was discovered in the season column a As the name implies, these are blood-feeding mites but are also present during later stages of decomposition b It is unusual for species in this genus to concentrate in huge proportions c It is unusual for species in this genus to concentrate in huge proportions. This genus is also very easy to confound with Poecilochirus or Parasitus species for a non-acarologist d Erroneous reports on websites; there is no evidence for Rostrozetes species being involved in animal decomposition or associated with carcases e It is unusual for species in this genus to concentrate in huge proportions. This genus is also very easy to confound with Poecilochirus or Parasitus species for a non-acarologist f Small animals: grey squirrel, fox squirrel, juvenile cottentail rabbit, cat, opossum; mites found on all carcases (Johnson 1975) Exp Appl Acarol (2009) 49:45–84 Exp Appl Acarol (2009) 49:45–84 73 fly maggot activity also renders the medium of the carcase alkaline, which is detrimental to the beetles. By reducing the amount of blow flies, the mites create a habitat more suitable for their phoretic hosts. However, this line of reasoning of a strictly mutual interaction is increasingly being questioned by acarologists. Poecilochirus mites might feed more on the carcase than on the blow fly eggs. Poecilochirus davydovae has now been recognized as a specialist predator feeding on the eggs of its beetle carrier, Nicrophorus vespilloides (Blackman 1997). Some mite species will end up at a carcase as incidentals, as species that use the corpse as a concentrated resource extension of their normal habitat; springtails (Collembola), spiders (Araneae), centipedes (Chilopoda), and wood lice (Isopoda) fall also in this category. However, mites as incidentals might be a minority group. Many mite species arrive at a carcase through phoresy on a necrophagous or necrophilous insect. The phoresy is often highly taxon specific. Many mite species arriving by phoresy are likely the product of evolutionary adaptation to a specialized food source and habitat, the opposite of incidental (Athias-Binche 1994; Perotti and Braig 2009b). But if mites are incidental, they might become the centre point of trace analysis in a forensic setting. Oligospecific infestations The importance of mites on carcases becomes even more pronounced under conditions of concealment or expedited dehydration, when the normal succession of arthropod waves is disrupted. Such situations often occur indoors. Carcases then decompose often completely under the action of a single or a few species of insects or mites. Insect species recorded in mono—or oligospecific infestations of human remains include the grey flesh fly Sarcophaga carnaria (=Musca carnaria; Sarcophagidae; Bergeret 1855), the brown house or false clothes moth Hofmannophila pseudospretella (=Borkhausenia pseudospretella; Oecophoridae; Forbes 1942), the corpse fly Hydrotaea capensis (Muscidae; Turchetto and Vanin 2004) or beetles. A case published by Schroeder et al. (2002) found that the leather or hide beetle Dermestes maculatus (Dermestidae) had almost skeletonised an indoor corpse in Germany within 5 months. A similar situation might have occurred involving the larder or bacon beetle D. lardarius in Denmark and the USA (Voigt 1965; Lord 1990). The first forensic case where mites have been used to estimate a post mortem interval involving a mummified corpse of a new-borne baby girl is also a case where one or two mite species were the only arthropods found on the corpse other than larvae of the grease moths Aglossa spp. (Pyralidae) (Brouardel 1879; Perotti 2009). The sprinkling or injection with lead arsenate of two human corpses found in the French Alps not only misled police dogs, but also prevented practically any insect infestation (Leclercq and Verstraeten 1992). The lead arsenate did not stop the bacterial decomposition. The bodies were mummified possibly through the effect of a dry and hot summer. With the exception of a very few fly larvae of miniscule size, the corpses carried only mites of the family Acaridae (=Tyroglyphidae), and even the mites were not in great numbers. In a more recent case reported from Germany, a child corpse found wrapped in plastic in a basement of a home was only associated with a mass occurrence of mites (Russell et al. 2004; OConnor 2009). Human corpses may be mosaics To assign a human corpse or any large carcase to a certain stage of decomposition might not be as straightforward as might be expected, especially, if the carcase is considered from an 123 74 Exp Appl Acarol (2009) 49:45–84 ecological point of view. Human body parts may be covered to varying degree with clothing that can have a drastic impact on decomposition. Exposed body parts like the face and hands might be skeletonised whereas clothed parts might still have most of the soft tissues in active or advanced stages of decay. Other parts of a carcase might develop adipocere or grave wax and enter a stage of mummification. This might be the case as much for an exposed body as for a body buried in a coffin. Particularly, woollen socks used to dress corpses in coffins have regularly delayed decomposition of soft tissue parts to a large degree. Clothed parts remained delayed in decomposition or preserved when exhumed after two or more years (Hunziker 1919). A human corpse sometimes might represent a mosaic of different stages of decomposition occurring simultaneously rather than a neat single stage. Often it is then just the biggest body part or the body part most advanced in the process of decomposition that determines the stage of decomposition represented in reports or in listings. The arthropod fauna present on such a corpse will reveal an increasing diversity the more carefully it is investigated. The more elaborate the clothing or other means of concealment, the stronger the impact on the decomposition process. The influence of clothing, wrapping and physical trauma such as knife wounds on the decomposition and arthropod succession has been studied in detail with pigs in central South Africa (Kelly 2006). The presence and absence of Acari during decomposition was recorded but not systematically analysed. A recent case of a child whose corpse had been wrapped in a pullover and plastic bag and hidden in a basement is illustrative (Russell et al. 2004). A water film formed on the inside of the plastic wrapping that generated a habitat characteristic of liquid decomposition at the transition between bloating stage and active decay. This liquid environment supported the mass occurrence of Myianoetus diadematus (Astigmata). At the same time, the rest of the body was at an advanced stage of decomposition characterised by the astigmatid mites Tyrophagus putrescentiae and Acarus immobilis; the corpse was probably 1–1.5 years post mortem. When the plastic bag was removed from the body, the M. diadematus colony collapsed through dehydration. Mites dominate in diversity and in numbers during the stages of butyric fermentation and dry decomposition. The low number of listings in the table for earlier stages of decomposition might be misleading. In the study of Johnson on small animals, all the mites were first recognised during the bloating stages but became very common during the dry decomposition stage (Johnson 1975). The mite presence spans four stages of decomposition. In a study with highly compromised chicken carcases with the flesh partially removed, Mesostigmata, Astigmata and Prostigmata were collected during the fresh stage (Arnaldos et al. 2004). Human mites Healthy humans will carry one or two species of symbiotic mites, Demodex brevis and D. folliculorum (Demodecidae, Prostigmata), the mites of sebaceous or fat glands and hair follicles (Desch 2009). These mites have been found on human corpses since their discovery in 1844 (Wilson 1844). Table 3 only gives exemplary references, for a more comprehensive account please see Perotti and Braig (2009a). Parasitic mites of humans do not feature during the fresh stage of Table 3, because humans have very few parasitic mites that are not incidental occurrences stemming from individual case reports. The best representative of a parasitic mite associated with humans is the scab mite Sarcoptes scabiei (Sarcoptidae, Astigmata). The sister species S. bovis of cows and S. equi of horses cause milker’s and cavalryman’s itch in humans during an abortive superficial infection. 123 Exp Appl Acarol (2009) 49:45–84 75 Cheyletiella blakei (Cheyletidae, Prostigmata) of cats, C. furmani and C. parasitivorax of rabbits and C. yasguri of dogs are mange mites, also known as walking dandruff, that might feed on epidermal keratin of humans and cause an abortive infection (Beesley 1998). Many chigger mites (Trombiculidae, Prostigmata) belonging to the genera Trombicula, Neotrombicula, Eutrombicula, Leptotrombicula and Ascoschoengastia may be encountered in the larval stage. These chiggers might feed on humans as an alternative host for a few days but are perhaps better regarded, like ticks and dermanyssid mites, as micropredators rather than as human parasites (Ashford and Crewe 2003). The species Eutrombicula belkini was central in linking a suspect to a murder scene in a case in California (Prichard et al. 1986; Turner 2009). Environment, microhabitats, size of carcase The impact of the habitat on the appearance of visible waves of Acari became evident in a comparative study using small pigs (around 9 kg) in three contrasting tropical habitats (Shalaby et al. 2000). Acari first became obvious 7–8 days post mortem in a mesophytic habitat, intermediate between dry and wet vegetation. At 11 days post mortem, Acari followed in the rain forest habitat of Oahu, Hawai’i (USA). Around 19–20 days post mortem, the pigs in the mesophytic and rain forest habitat experienced a second wave of mites; and pigs in an arid, xerophytic habitat received their first wave of mites. Studies of the insects associated with small carcases have been characterised by dramatic variations in the carrion-feeding fauna (Blackith and Blackith 1990). Even small variations in the size of the carcase may have an influence on the stage at which mites are obvious. For very small pigs of 8.4 kg, nymphs and adults of Acaridae (Astigmata) and Macrochelidae (Mesostigmata) and adults of Liacaridae (Oribatida) were dominant during the postdecay stage, 12–16 days post mortem, whereas the same mite population occurred during the remains stage, 14–30? days post mortem, for a pig carcase of 15.1 kg (Hewadikaram and Goff 1991). The seasons can have a huge impact on the stage of decomposition at which mites become obvious. In a study in a farmland area in the north of Spain using pigs exposed to the sun, mites became obvious at the fresh stage during winter, at the bloating stage during spring, at the active decomposition stage during autumn, and remained absent even at the advanced stage of decomposition during summer (Castillo Miralbes 2002). However, in experiments with chicken carcases with the flesh partially removed and the viscera present showed the highest numbers of mites (687) during summer and advanced decomposition followed by spring (216); winter had 190 mites during the earlier stage of decomposition and autumn showed overall the lowest numbers (Arnaldos et al. 2004). The chicken carcases were put in an agricultural field around Murcia in southeastern Spain. The impact of the season on the abundance of mites on a carcase also becomes evident if the numbers of mites are put in relation to other major sarcosaprophagous arthropods. The percentual contribution of mites to the fauna on the chicken carcases can almost be as high as that of flies during the summer, and during winter still much higher than that of beetles: spring: 42% Diptera, 33% Hymenoptera, 9% Collembola, 5% Acari, 3% Coleoptera; summer: 29% Hymenoptera, 22% Diptera, 21% Acari, 14% Collembola, 5% Coleoptera; autumn: 55% Collembola, 37% Diptera, 3% Hymenoptera, 2% Coleoptera, 1% Acari; winter: 41% Diptera, 39% Collembola, 8% Acari, 2% Coleoptera, Hymenoptera and Psocoptera, each (major constituents only) (Arnaldos Sanabria 2000; Goff et al. 2004). 123 76 Exp Appl Acarol (2009) 49:45–84 The pig study also showed that carcases exposed to the sun during autumn contained mites at the active or advanced stage of decomposition, whereas carcases kept at the same time in a shadowed environment 300 m away already had mites at the bloating stage. The differences might be explained to a great extent by the scotophilic or heliophilic behaviour of the insects carrying the mites. Both, shadow and lower temperatures facilitate early mite colonisation of carcases in the pig experiments. The fact that many mite species are photonegative can make the collection of mites during daylight or in direct sunlight difficult and unrepresentative for the actual diversity and abundance present. The seasons also have some influence on the families of mites colonising the carcase. Hard ticks (Ixodidae) were only found during spring at the bloated stage and at active decomposition in the shadow, and during winter at active decomposition in the sun. Since ticks are obligate parasites of living animals, the presence of ticks might reflect the activity of scavengers at that time (Castillo Miralbes 2002). The study with chickens confirms the presence of hard ticks only during spring time (Arnaldos et al. 2004). A comprehensive study on the influence of shade and sun exposure with pigs was performed in Edmonton, Canada (Anderson et al. 2002). Careful records on the presence or absence of mites during decomposition were kept but mites were not systematically differentiated. Mite dispersal The importance of phoresy for the introduction of mites to carcases has repeatedly been emphasised; for review, see Perotti et al. (2009a). Often overlooked is the fact that these mites also have to leave the carcase again at a certain time. Skin beetles (Trogidae) can become so heavily overloaded that their mites also infest and cover larval stages, which have no functional role in phoresy. The infestation can become so severe that the beetles end up dead in and around the carcase. This has also been observed for skin beetles on pig carcases and beetles in general on dog carcases (Reed 1958; Gill 2005). Macrocheles species go to their beetle species. Parasitus and Poecilochirus species jump on everything that moves and easily saturate the phoretic host. Details of mite-host associations can be found in Perotti and Braig (2009b). The end of a wave of either mites or their insect carriers might be judged by the level of mite infestation on a particular carrier. Another aspect of dispersal is the analysis of mites that were already present before death. Very few studies have addressed this point. One study on pigs in Nigeria observed that the ticks present naturally on the pig left the pig to find a new host as the bloated stage approached (Iloba and Fawole 2006). Humans carry mites in hair follicles and skin pores but also on their clothing (Desch 2009; Perotti and Braig 2009a). The diversity of mites found in buildings and homes might gain forensic importance (Frost et al. 2009; Solarz 2009; Colloff 2009). Using furred or feathered animals in forensic experiments as substitutes for human bodies poses some problems for the investigation of mites. A study of mites on rat species showed that many parasitic mite species present in the fur during life are still recovered from the dead animals (Ramsay and Paterson 1977). Even feather mites were found on the rats. The diversity of known mite species associated with fur and feathers is huge and might represent only 20% of the actual number. For example, there are more than 2,000 feather mite species described belonging to 44 genera and 33 families. Only pigs, elephants, rhinoceroses, mole rats, whales and hippopotamuses share naturally the nakedness 123 Exp Appl Acarol (2009) 49:45–84 77 with humans. Mexican hairless dogs and sphinx cats might be alternatives but have no advantage over pigs. Unfortunately, the only decomposition study on elephants did not consider mites (Coe 1978). The soil below Mites might be the most abundant soil invertebrates beneath a carcase (Anderson and VanLaerhoven 1996). Bornemissza (1957) studied the impact of decomposing guinea pigs on the natural soil fauna beneath the carcases in Perth, Western Australia. He graphically showed that on the soil surface and in the soil to a depth of 15 cm, the natural mite fauna together with most other arthropod taxa seem to mainly disappear 5–6 days into the decomposition process and reappear some 3 months later. The complete absence of oribatid mites or subterranean springtails such as Onychiuris and Tullbergia spp. indicated that the reduction of the typical soil fauna was very severe. It was greatest under the oral and anal parts of the carcase. These graphs and this information have been widely cited in the forensic entomological literature suggesting that the fauna beneath a carcase might be highly impoverished during most of the decomposition process and therefore of little forensic interest. This, however, might actually have been exceptional and should not be generalised. Bornemissza, citing Kühnelt (1950), also states that in Europe mites are only present during the final stages of decomposition. We don’t see any evidence for such assertions. However, we have no doubt that soil mites under carcases will display geographical behavioural variation, caused by climatic or edaphic factors (Dadour and Harvey 2008). Reed (1958) in a study with dogs in Tennessee described that soil samples taken beside carcases teemed with mites. At various times mites were piled in layers several individuals thick on the putrefactive substance under carcases. They were most abundant during warm and hot weather, but during the winter a few mites could generally be found under each carcase. In a study with cats on the island of Oahu, Hawai’i, Goff not only demonstrated large quantities of mites but also showed that changes in mesostigmatid populations (Macrochelidae, Parasitidae, Uropodidae and Pachylaelapidae) in samples of soil and litter removed from under the carcases could be correlated with post mortem intervals (Goff 1989). Goff reported on a homicide case where soil was found in the hood of a jacket that had been associated with the skull of a child of approximately 30 months of age recovered from a shallow grave on a narrow ledge on the side of Koko Head Crater on Oahu (Goff 1991). This is the third case in a comparative study by Goff of human decomposition ranging from 8 to 53 days post mortem reported earlier (Goff and Odom 1987). The soil exhibited a rich diversity of mite taxa that had previously been found on and under pig and cat carcases. The taxa are listed in Table 3. Although the acarine fauna considered in this case was not by itself definitive of a specific post-mortem interval, it served to provide valuable supporting data for the refining of the estimate toward the lower end of the window defined by the insects collected from the corpse (Goff 1991). The insect data suggested a period between 51 and 76 days. Presence of only adults of two species of Macrochelidae was consistent with an interval of 22–60 days. Presence of numbers of T. putrescentiae was characteristic of a time period greater than 48 days. Other mite species present were not definitive of any time period for this case. There was a total of 97 mites/10 cm3 of soil for this sample, a number corresponding to an interval of 48–52 days in decomposition studies previously conducted. Based on the estimated post mortem interval, the authorities requestioned the father of the child. In his subsequent 123 78 Exp Appl Acarol (2009) 49:45–84 confession he put the time of death at the 53rd day prior to the collection of the samples (Goff 1991). In a study with bank vole carcases in a wooded park in Poland with acid soil, it was noticed that carcases left on the surface experienced mite infestations during the initial stages of decomposition and during the final residual stages with little mite participation during active decomposition. However, when the carcases were buried in a 25–30 cm deep hole, mites dominated during active decomposition and residual stages but not during the initial process (Nabagło 1973). The soil of a large wooded area in Massachusetts during summer harboured mites of the families Acaridae (Asigmata), Digamasellidae, Laelapidae, Uropodidae (Mesostigmata), and Nothridae (Oribatida) under turtle carcases as well as in control samples (Abell et al. 1982). Northridae were found in very small numbers and Laelapidae in large numbers also on the turtle carcases themselves. The forest consisted of a mixture of deciduous trees primarily made up of red oak and red maple with some American beech and white pine. The soil beneath the carcases contained in addition the following families: Ceratozetidae (Oribatida), Diplogyniidae (Mesostigmata) and Rhagidiidae (Prostigmata), while soil far from the carcases also contained the families Galumnidae, Hypochthoniidae (Oribatida) and Phytoseiidae (Mesostigmata). The dominant family on the turtles and in the soil beneath exposed carrion was Laelapidae. Payne et al. (1968) compared the mite families on surface exposed baby pigs and baby pigs in burial pits at depths varying from 50 to 100 cm. Twenty-six of 48 arthropod species were not implicated in above-ground carrion succession, but were found only on buried pigs; among these were the mite families Uropodidae and Acaridae. Mummies might harbour mites belonging to the Tarsonemidae (Prostigmata) and/or mites in general that are associated with a practice of food storage, food gifts or the use of raw cotton to wrap the corpse, oribatid mites that often originate from soil contaminations, or mites that might be derived from plant material in general or leaves of coca added to the corpse (Leles de Souza et al. 2006; Mendonça de Souza et al. 2008; Baker 2009). Coprolites and faeces Corpses also come with faeces, and faeces attract mites. A great diversity of mites has been collected from inside human mummies (Baker 2009). Practically no work has been done on the mites attracted to relatively fresh faeces of human corpses. It seems that more acarological information is available on coprolites of human and animal mummies (Radovsky 1970; Kliks 1988; de Candanedo Guerra et al. 2003) or 6,500 year-old Demodex mites in regurgitated pellets of raptors (Fugassa et al. 2007). Radovsky identified deutonymphs of Myianoetus nr dionychus and Anoetostoma oudemansi (Histiostomatidae, Astigmata) and an acarid tritonymph in a human coprolite (Radovsky 1970). Mass occurrence of M. diadematus, a species related to M. nr dionychus, was recently reported from the corpse of a human baby wrapped in a plastic bag (Russell et al. 2004). The histiostomatid and acarid mites found there might have been attracted by the fresh faeces; however, mites of these two families might also have been ingested with food and passed in the faeces, something that happens unnoticed but perhaps frequently in most human cultures (Radovsky 1970). Acknowledgments The authors appreciate the funding of research on forensic acarology by the Leverhulme Trust. Additional information was kindly provided by M. Lee Goff, Paola Magni, Marta I. SaloñaBordas and Francis D. Feugang Youmessi. The authors like to thank Mariló Moraza and Barry M. OConnor for advice and reviewing an earlier version of the manuscript. 123 Exp Appl Acarol (2009) 49:45–84 79 References Abell DH, Wasti SS, Hartmann GC (1982) Saprophagous arthropod fauna associated with turtle carrion. Appl Entomol Zool 17:301–307 Anderson GS, Vanlaerhoven SL (1996) Initial studies on insect succession on carrion in southwestern British Columbia. J Forensic Sci 41:617–625 Anderson GS, Hobischak N, Samborski C et al (2002) Insect succession on carrion in the Edmonton, Alberta, region of Canada Technical Report TR-04-2002. Canadian Police Research Centre, Ottawa (Ontario), Canada Arnaldos MI, Romera E, Presa JJ et al (2004) Studies on seasonal arthropod succession on carrion in the southeastern Iberian Peninsula. Int J Legal Med 118:197–205 Arnaldos Sanabria MI (2000) Estudio de la fauna sarcosapprófaga de la Región de Murcia. Su aplicación a la mediciona legal [Studies on the sarcosaphrophagous fauna in the Region of Murcia; its application in legal medicine]. Departamento de Biologı́a Animal. Universidad de Murcia, Murcia Athias-Binche F (1994) La Phorésie chez les Acariens—Aspects Adaptatifs et Evolutifs [Phoresy in acarina—adaptive and evolutionary aspects]. Editions du Castillet, Perpignan Ashford RW, Crewe W (2003) The parasites of Homo sapiens. An annotated checklist of the protozoa, helminths and arthropods for which we are home. Taylor & Francis, London Avila FW, Goff ML (1998) Arthropod succession patterns onto burnt carrion in two contrasting habitats in the Hawaiian islands. J Forensic Sci 43:581–586 Baker AS (1990) Two new species of Lardoglyphus Oudemans (Acari: Lardoglyphidae) found in the gut contents of human mummies. J Stored Prod Res 26:139–147 Baker AS (2009) Acari in archaeology. Exp Appl Acarol 49. doi:10.1007/s10493-009-9271-1 Beesley WN (1998) Scabies and other mite infestations. In: Palmer SR, Lord Soulsby EJL, Simpson DIH (eds) Zoonoses. Oxford University Press, Oxford, pp 859–872 Behan-Pelletier V, Bissett B (1994) Oribatida of Canadian peatlands. Mem Entomol Soc Can 169:73–88 Bergeret M (1855) Infanticide. Momification naturelle du cadavre. Découverte du cadavre d’un enfant nouveau-né dans une cheminée où il s’était momifié. Détermination de l’époque de la naissance par la présence de nymphes et de larves d’insectes dans le cadavre et par l’étude de leurs métamorphoses [Infanticide. Natural mummification of the corpse. A corpse of a new-born child discovered in a chimney where it had been mummified. Determination of the time of the birth by the presence of nymphs and larvae of insects in the corpse and by the study of their metamorphoses]. Ann Hyg Publ Méd Lég 4(série):442–452 Bianchini G (1929) Contributo pratico e sperimentale allo studio della fauna cadaverica [An applied and experimental contribution to the study of the cadervous fauna]. Atti Accad Fisiocrit Siena 4(serie 10):97–106 Blackith RE, Blackith RM (1990) Insect infestations of small corpses. J Nat Hist 24:699–709 Blackman S (1997) Experimental evidence that the mite Poecilochirus davydovae (Mesostigmata: Parasitidae) eats the eggs of its beetle host. J Zool 242:63–67 Bornemissza GF (1957) An analysis of arthropod succession in carrion and the effect of its decomposition on the soil fauna. Aust J Zool 5:1–12 Bourel B, Tournel G, Hédouin V et al (2004) Entomofauna of buried bodies in northern France. Int J Legal Med 118:215–220 Braack LEO (1986) Arthropods associated with carcasses in the northern Kruger national park. S Afr J Wildl Res 16:91–98 Braack LEO (1987) Community dynamics of carrion-attendant arthropods in tropical African woodland. Oecologia 72:402–409 Bregetova NG, Koroleva EV (1960) The macrochelid mites (Gamasoidea, Macrochelidae) in the USSR. Parazitol Sb 19:32–154 Brouardel P (1879) De la détermination de l’époque de la naissance et de la mort d’un nouveau-née, faite à l’aide de la présence des acares et des chenilles d’aglosses dans cadavre momifié [Determination of the time of birth and of death of a new-born child, made using the presence of mites and Aglossa caterpillars on the mummified corpse]. Ann Hyg Publ Méd Lég 2(série 3):153–158 Brown JM, Wilson DS (1994) Poecilochirus carabi: behavioral and life-history adaptations to different hosts and the consequences of geographical shifts in host communities. In: Houck MA (ed) Mites. Ecological and evolutionary analyses of life history patterns. Chapman and Hall, New York, pp 1–22 Castillo Miralbes M (2002) Estudio de la entomofauna asociada a cadáveres en el Alto Aragón (España) [Study of the entomofauna associated with corpses in the region of Alto Aragón (Spain)]. Sociedad Entomológica Aragonesa, Zaragoza 123 80 Exp Appl Acarol (2009) 49:45–84 Chapman RF, Sankey JHP (1955) The larger invertebrate fauna of three rabbit carcasses. J Anim Ecol 24:395–402 Coe M (1978) The decomposition of elephant carcasses in the Tsavo (East) National Park, Kenya. J Arid Environ 1:71–86 Collins M (1970) Studies on the decomposition of carrion and its relationship with its surrounding ecosystem. PhD Thesis, Department of Zoology, University of Reading, Reading, England Colloff MJ (2009) Dust mites. Springer, Dordrecht Cornaby BW (1974) Carrion reduction by animals in contrasting tropical habitats. Biotropica 6:51–63 Dadour IR, Harvey ML (2008) The role of invertebrates in terrestrial decomposition: forensic applications. In: Tibbett M, Carter DC (eds) Soil analysis in forensic taphonomy. CRC Press, Boca Raton, pp 109–122 Dahl F (1896) Vergleichende Untersuchungen über die Lebensweise wirbelloser Aasfresser [Comperative studies on the ecology of invertebrate carrion feeders]. Sitzungsb Königl Preuss Akad Wiss Berlin 1:17–30 Davis JB, Goff ML (2000) Decomposition patterns in terrestrial and interdidal habitats on Oahu Island and Coconut Island, Hawaii. J Forensic Sci 45:836–842 de Candanedo Guerra RdMSN, Gazeta GS, Amorim M et al (2003) Ecological analysis of Acari recovered from coprolites from archaeological site of Northeast Brazil. Mem Inst Oswaldo Cruz 98(Suppl. 1):181–190 De Jong GD, Chadwick JW (1999) Decomposition and arthropod succession on exposed rabbit carrion during summer at high altitudes in Colorado, USA. J Med Entomol 36:833–845 De Jong GD, Hoback WW (2006) Effect of investigator disturbance in experimental forensic entomology: succession and community composition. Med Vet Entomol 20:248–258 Desch CE (2009) Human hair follicle mites and forensic acarology. Exp Appl Acarol 49. doi: 10.1007/s10493-009-9272-0 Early M, Goff ML (1986) Arthropod succession patterns in exposed carrion on the island of O’ahu, Hawaiian islands, USA. J Med Entomol 23:520–531 Easton AM, Smith KGV (1970) The entomology of the cadaver. Med Sci Law 10:208–215 Feugang Youmessi FD, Djiéto-Lordon C, Gaudry E et al (2008) Contribution to the research of the entomological indicators of corpse dating: case of Rattus rattus (Linnaeus, var WISTAR) in Yaounde (Cameroon) EAFE Meeting 2008, Kolymbari, Greece Forbes G (1942) The brown house moth as an agent in the destruction of mummified human remains. Police J Lond 15:141–148 Fourman KL (1936) Kleintierwelt, Kleinklima, und Mikroklima in Beziehung zur Kennzeichnung des Forstlichen Standorts und der Bestandsabfallzersetzung auf bodenbiologischer Grundlage [Microfauna, local climate, and microclimate in relationship with the characterisation of forest location and decomposition of forest waste on a soil-biological basis]. Mitt Forstwirt Forstwiss 7:596–615 Frost CL, Amendt J, Braig HR, Perotti MA (2009) Indoor arthropods of forensic importance. In: Amendt J, Goff ML, Campobasso CP et al (eds) Current concepts in forensic entomology. Springer, Dordrecht Fugassa MH, Sardella NH, Denegri GM (2007) Paleoparasitological analysis of a raptor pellet from Southern Patagonia. J Parasitol 93:421–422 Fuller ME (1934) The insect inhabitants of carrion: a study in animal ecology. Council of Science and Industry Research in Australia, Canberra Gaudry E (2002) Eight squadrons for one target: the fauna of cadaver described by P. Mégnin Proceedings of the First European Forensic Entomology Seminar, Rosny sous Bois, France, pp 23–28 Gill GJ (2005) Decomposition and arthropod succession on above ground pig carrion in rural Manitoba Technical Report TR-06-2005. Canadian Police Research Centre, Ottawa (Ontario) Gmeiner F (1908) Demodex folliculorum des Menschen und der Tiere [Demodex folliculorum of humans and animals]. Arch Dermatol Syph 92:25–96 Goff ML, Odom CB (1987) Forensic entomology in the Hawaiian Islands: three case studies. Am J Forensic Med Pathol 8:45–50 Goff ML (1989) Gamasid mites as potential indicators of postmortem interval. In: Channabasavanna GP, Viraktamath CA (eds) Progress in Acarology, vol 1. Oxford & IBH Publishing, New Delhi, pp 443–450 Goff ML (1991) Use of acari in establishing a postmortem interval in a homicide case on the island of Oahu, Hawaii. In: Dusbábek E, Bukva V (eds) Modern Acarology, vol 1. SPB Academic Publishing, The Hague, pp 439–442 Goff ML (1993) Estimation of postmortem interval using arthropod development and successional patterns. Forensic Sci Rev 5:81–94 123 Exp Appl Acarol (2009) 49:45–84 81 Goff ML, Garcı́a Garcı́a MD, Arnaldos Sanabria MI (2004) Entomologı́a cadavérica: Fundamentos y aplicación. Referencia a la entomologı́a española [Forensic entomology: basics and applications. A reference to Spanish entomology]. In: Gisbert Calabuig JA, Villanueva Cañadas E et al (eds) Tratado de Medicina Legal y Toxicologı́a [Treatise on legal medicine and toxicology]. Masson, Barcelona, pp 253–273 Goff ML (2009) Early post-mortem changes and stages of decomposition in exposed cadavers. Exp Appl Acarol 49. doi:10.1007/s10493-009-9284-9 Graells M (1886) Entomologia judicial [Forensic entomology]. Rev Progr Cienc Exact Fı́s Nat Madrid 21:458–471 Grassberger M, Frank C (2004) Initial study of arthropod succession on pig carrion in a central European urban habitat. J Med Entomol 41:511–523 Gwiazdowicz DJ, Klemt J (2004) Mesostigmatic mites (Acari, Gamasida) in selected microhabitats of the Biebrza National Park (NE Poland). Biol Lett 41:11–19 Halliday RB (2000) The Australian species of Macrocheles (Acarina : Mesostigmata). Invertebr Taxon 14:273–326 Haskell NH, Hall RD, Cervenka VJ (1997) On the body: insect’s life stage presence and their postmortem artefacts. In: Hagland WD, Sorg MH et al (eds) Forensic taphonomy–the post mortem fate of human remains. CRC Press, Boca Raton, pp 415–467 Hewadikaram KA, Goff ML (1991) Effect of carcass size on rate of decomposition and arthropod succession patterns. Am J Forensic Med Pathol 12:235–240 Hidalgo-Argüello MR, Dı́ez Baños N, Fregeneda Grandes J et al (2003) Parasitological analysis of Leonese royalty from Collegiate-Basilica of St. Isidoro, Léon (Spain): Helminths, protozoa, and mites. J Parasitol 89:738–743 Hobischak NR, Anderson GS (2002) Time of submergence using aquatic invertebrate succession and decompositional changes. J Forensic Sci 47:143–151 Horenstein MB, Arnaldos MI, Rosso B et al (2005) Estudio preliminar de la comunidad sarcosaprófaga en Córdoba (Argentina): aplicación a la entomologı́a forense [Preliminary study of the sarcosaprophytic community in Cordoba (Argentina): applied to forensic entomology]. An Biol 27:191–201 Hunziker H (1919) Über die Befunde bei Leichenausgrabungen auf den Kirchhöfen Basels. Unter besonderer Berücksichtung der Fauna und Flora der Gräber [About the findings during excavations of corpses on the cemeteries of Basel, especially of the fauna and flora of graves]. Frankf Z Pathol 22:147–207 Hyatt KH, Emberson RM (1988) A review of the Macrochelidae (Acari: Mestostigmata) of the British Isles. Bull Br Mus (Natl Hist) Zool 54:63–125 Iloba BN, Fawole SO (2006) Comparative study of arthropod fauna on exposed arrions across the vertebrate classes. Int J Biomed Health Sci 2:51–65 Johnson MD (1975) Seasonal and microseral variation in the insect populations on carrion. Am Midl Nat 93:79–90 Johnston W, Villeneuve G (1897) On the medico-legal application of entomology. Montr Med J 26:81–89 Kelly JA (2006) The influence of clothing, wrapping and physical trauma on carcass decomposition and arthropod succession in central South Africa. PhD Thesis, Department of Zoology and Entomology, University of the Free State, Bloemfontein, South Africa Kliks MM (1988) Paleoparasitological analyses of fecal material from Amerindian (or New World) mummies: evaluation of saprophytic arthropod remains. Paleopathol Newsl 64:7–11 Kneidel KA (1984) Competition and disturbance in communities of carrion-breeding Diptera. J Anim Ecol 53:849–865 Krantz GW, Whitaker JO Jr (1988) Mites of the genus Macrocheles (Acari: Macrochelidae) associated with small mammals in North America. Acarologia 29:225–259 Krantz GW, Platnick NI (1995) On Brucharachne, the spider that wasn’t (Arachnida, Acari, Dermanyssoidea). Am Mus Novit 3151:1–8 Kühnelt W (1950) Bodenbiologie [Soil biology]. Herold, Vienna Lecha-Marzo A (1917) Tratado de autopsias y embalsamamientos [Treatise on autopsy and embalming]. Los Progresos de la Clı́nica, Madrid, pp 79–90 Leclercq M (1978) Entomologie et Médecine Légale: Datation de la Mort [Entomology and forensic medicine: dating the time of death]. Masson, Paris Leclercq M (2002) L’entomologie légale en Belgique depuis 1947 [Forensic entomology in Belgium since 1947] Proceedings of the First European Forensic Entomology Seminar, Rosny sous Bois, France, pp 8–12 Leclercq M, Verstraeten C (1988a) Entomologie et médicine légale. Datation de la mort: insectes et autres arthropodes trouvés sur les cadavres humains [Entomology and forensic medicine, determination of the 123 82 Exp Appl Acarol (2009) 49:45–84 time of death: insects and other arthropods on human cadavers]. Bull Ann Soc R Belge Entomol 124:311–317 Leclercq M, Verstraeten C (1988b) Entomologie et médecine légale. Datation de la mort. Acariens trouvés sur des cadavres humains [Entomology and forensic medicine. Determination of the time of death. Acari found on human cadavers]. Bull Ann Soc R Belge Entomol 124:195–200 Leclercq M, Verstraeten C (1992) Eboueurs entomologiques bénévoles dans les écosystèmes terrestres: observation inédite [Voluntary entomological street sweepers in the terrestrial ecosystems: a new observation]. Notes Faun Gembloux 25:17–22 Leclercq M, Verstraeten C (1993) Entomologie et médecine légale. L’entomofaune des cadavres humains: sa succession par son interprétation, ses résultats, ses perspectives [Entomology and forensic medicine. The entomofauna of human corpses: its succession and interpretation, its results, its prospects]. J Med Leg Droit Med 36:205–222 Leclerq M (1969) Entomological parasitology: the relations between entomology and the medical sciences. Pergamon, Oxford Leles de Souza D, de Maria Seabra Nogueira de Candanedo Guerra R, Mendonça de Souza S et al (2006) Acari found in a mummy bundle from the Chillon River Valley, Peru. Paleopathol Newsl 136:11–16 Lichtenstein J, Moitessier A, Jaumes A (1885) Un nouveau cas d’application de l’entomologie à la médecine légale [A new case of the application of entomology in legal medicine]. Ann Hyg Publ Méd Lég 13(série 3):121–127 Lord WD, Burger JF (1984a) Arthropods associated with harbor seal (Phoca vitulina) carcasses stranded on islands along the New England Coast. Int J Entomol 26:282–285 Lord WD, Burger JF (1984b) Arthropods associated with herring gull (Larus argentatus) and great blackbacked gull (Larus marinus) carrion on islands in the Gulf of Maine. Environ Entomol 13:1261–1268 Lord WD (1990) Case histories of the use of insects in investigations. In: Catts EP, Haskell NH (eds) Entomology & death: a procedural guide. Joyce’s Print Shop, Clemson, pp 9–37 Mašán P (1993) Mites (Acarina) associated with species of Trox (Coleoptera: Scarabaeidae). Eur J Entomol 90:359–364 Magni P, Ghizzoni O, Linarello P et al (2008) The man in the farm house—effective support of entomotoxicological examinations to identify causes of death EAFE Meeting 2008, Kolymbari, Greece Mégnin P (1887) La faune des tombeaux [The fauna of graves]. C R Hebd Acad Sci 105:948–951 Mégnin P (1894) La Faune des Cadavres. Application de l’Entomologie à la Médecine Légale [The fauna of corpses. Application of entomology to forensic medicine]. G. Masson and Gauthier-Villars et Fils, Paris Mégnin P (1895) La faune des cadavres [The fauna of carcasses]. Ann Hyg Publ Méd Lég série 3(33):64–67 Mégnin P (1898) Les parasites de la mort. Une cause peu connue de la momification des cadavres [Parasites of death. A little known cause of the mummification of the corpses]. Arch Parasitol 1:39–43 Mendonça de Souza SMF, Reinhard KJ, Lessa A (2008) Cranial deformation as the cause of death for a child from the Chillon River Valley, Peru. Chungará 40:41–53 Merritt RW, Snider R, de Jong JL et al (2007) Collembola of the grave: a cold case history involving arthropods 28 years after death. J Forensic Sci 52:1359–1361 Michelsen V (1983) Thyreophora anthropophaga, an extinct bone skipper rediscovered in Kashmir, India (Diptera, Piophilidae, Thyreophorina). Entomol Scand 14:411–414 Motter MG (1898) A contribution to the study of the fauna of the grave. A study of on hundred and fifty disinterments, with some additional observations. J N Y Entomol Soc 6:201–233? Nabagło L (1973) Participation of invertebrates in decomposition of rodent carcasses in forest ecosystems. Ekol Polska 21:251–270 OConnor BM (2009) Astigmatid mites (Acari: Sarcoptiformes) of forensic interest. Exp Appl Acarol 49. doi:10.1007/s10493-009-9270-2 Parker JC, Holliman RB (1971) Observations on parasites of gray squirrels during the 1968 emigration in North Carolina. J Mammal 52:437–441 Payne JA (1965) A summer carrion study of the baby pig Sus scrofa Linnaeus. Ecology 46:592–602 Payne JA, Crossley DAJ (1966) Animal species associated with pig carrion Oak Ridge. National Laboratory Technical Memorandum, Oak Ridge Payne JA, King EW, Beinhart G (1968) Arthropod succession and decomposition of buried pigs. Nature 219:1180–1181 Payne JA, King EW (1972) Insect succession and decomposition of pig carcasses in water. J Georgia Entomol Soc 34:153–162 Pérez SP, Duque P, Wolff M (2005) Successional behavior and occurrence matrix of carrion-associated arthropods in the urban area of Medellı́n, Colombia. J Forensic Sci 50:448–454 123 Exp Appl Acarol (2009) 49:45–84 83 Perotti MA (2009) Mégnin re-analysed: the case of the newborn baby girl, Paris, 1878. Exp Appl Acarol 49. doi:10.1007/s10493-009-9279-6 Perotti MA, Braig HR (2009a) Acarology in criminolegal investigations: the human acarofauna during life and death. In: Byrd JH, Castner JL (eds) Forensic entomology: the utility of arthropods in legal investigations. Taylor & Francis, Boca Raton, pp 637–649 Perotti MA, Braig HR (2009b) Phoretic mites associated with animal and human decomposition. Exp Appl Acarol 49. doi:10.1007/s10493-009-9280-0 Perotti MA, Braig HR, Goff ML (2009a) Phoretic mites and carcasses. In: Amendt J, Goff ML, Campobasso CP et al (eds) Current concepts in forensic entomology. Springer, Dordrecht Perotti MA, Goff ML, Baker AS et al (2009b) Forensic acarology, an introduction. Exp Appl Acarol 49 (in press) Pont AC, Matile L (1980) Découverte de quelques insectes de J.-P. Mégnin; identité d’Ophyra cadaverina Mégnin (1894) (Diptera, Muscidae) [Discovery of several insects of J.-P. Mégnin; identity of Ophyra cadaverina Mégnin (1894) (Diptera, Muscidae)]. Bull Soc entomol France 85:41–43 Porta CF (1929) Contributo allo studio dei fenomeni cadaverici: L’azione della microfauna cadaverica terrestre nella decomposizione del cadavere [Contribution to the study of cadaveral phenomina: the behaviour of the terrestrial microfauna of cadavers during the decomposition of cadavers]. Arch Antropol Crim Psich Med Leg Sci Aff 59:1–55 Prichard JG, Kossoris PD, Leibovitch RA et al (1986) Implications of trombiculid mite bites: reports of a case and submission of evidence in a murder trial. J Forensic Sci 31:301–306 Proctor HC (2009) Can freshwater mites act as forensic tools? Exp Appl Acarol 49. doi: 10.1007/s10493-009-9273-z Putman RJ (1978) The role of carrion-frequenting arthropods in the decay process. Ecol Entomol 3:133–139 Radovsky FJ (1970) Mites associated with coprolites and mummified human remains in Nevada. Contr Univ Calif Archaeol Res Facility 10:186–190 Ramsay GW, Paterson SE (1977) Mites (Acari) from Rattus species on Raoul Island. N Z J Zool 4:389–392 Reed HB Jr (1958) A study of dog carcass communities in Tennessee, with special reference to the insects. Am Midl Nat 59:213–245 Richards EN, Goff ML (1997) Arthropod succession on exposed carrion in three contrasting tropical habitats on Hawaii Island, Hawaii. J Med Entomol 34:328–339 Rı́os T (1902a) Los insectos y la putrefacción de los cadáveres [Insects and the decomposition of corpses] (I-II). Clı́n Mod Rev Med Cirug 1:74–80 Rı́os T (1902b) Los insectos y la putrefacción de los cadáveres [Insects and the decomposition of corpses] (III-VI). Clı́n Mod Rev Med Cirug 1:171–180 Rives DV, Barnes HJ (1988) Pseudoparasitism of broiler chicks by mites of the family Uropodidae, genus Fuscuropoda. Avian Dis 32:567–569 Russell DJ, Schulz MM, OConnor BM (2004) Mass occurence of astigmatid mites on human remains. Abh Ber Naturkundemus Görlitz 76:51–56 Samšiňák K (1960) Über einige myrmekophile Milben aus der Familie Acaridae [On some myrmecophylic mites in the family Acaridae]. Čas Česk Spol Entomol 57:185–192 Schnell e Schühli G, de Carvalho CJB, Wiegmann BM (2004) Regarding the taxonomic status of Ophyra Robineau-Desvoidy (Diptera: Muscidae): a molecular approach. Zootaxa 712:1–12 Schnell e Schühli G, de Carvalho CJB, Wiegmann BM (2007) Molecular phylogenetics of the Muscidae (Diptera: Calyptratae): new ideas in a congruence context. Invertebr Syst 21:263–278 Schoenly KG, Shahid SA, Haskell NH et al (2005) Does carcass enrichment alter community structure of predaceous and parasitic arthropods? A second test of the arthropod saturation hypothesis at the anthropology resaerch facility in Knoxville, Tennessee. J Forensic Sci 50:134–142 Schönborn W (1963) Vergleichende zoozönotische Untersuchungen an Exkrementen, Kadavern, Hutpilzen und Vogelnestern [Comparative zoocenotic investigations on excrements, carcasses, mushrooms and bird nests]. Biologisches Zentralbl 82:165–184 Schroeder H, Klotzbach H, Oesterhelweg L et al (2002) Larder beetles (Coleoptera, Dermestidae) as an accelerating factor for decomposition of a human corpse. Forensic Sci Int 127:231–236 Shalaby OA, deCarvalho LML, Goff ML (2000) Comparison of patterns of decomposition in a hanging carcass and a carcass in contact with soil in a xerophytic habitat on the island of Oahu, Hawaii. J Forensic Sci 45:1267–1273 Smith KGV (1973) Forensic entomology. In: Smith KGV (ed) Insects and other arthropods of medical importance. British Museum (Natural History). London, UK, pp 483–486 Smith KGV (1975) The faunal succession of insects and other invertebrates on a dead fox. Entomol Gaz 26:277–287 Smith KGV (1986) A manual of forensic entomology. British Museum (Natural History), London 123 84 Exp Appl Acarol (2009) 49:45–84 Solarz K (2009) Indoor and dust mites. Exp Appl Acarol 49. doi:10.1007/s10493-009-9292-9 Strauch C (1912) Die Fauna der Leichen [Fauna of corpses]. Vierteljahrsschr gerichtl Med öffentl Sanitätsw 43:44–49 Strauch C (1928) Beiträge zur natürlichen Mumifikation menschlicher Leichen [Contribution to the natural mummification of human corpses]. Dtsch Z gesamte gerichtl Med 12:259–269 Tantawi TI, El-Kady EM, Greenberg B et al (1996) Arhropod succession on exposed rabbit carrion in Aexandria, Egypt. J Med Entomol 33:566–580 Turchetto M, Vanin S (2004) Forensic evaluations on a crime case with monospecific necrophagous fly population infected by two parasitoid species. Aggrawal’s Internet J Forensic Med Toxicol 5:12–18 Turner B (2009) Forensic entomology: a template for forensic Acarology? Exp Appl Acarol 49. doi: 10.1007/s10493-009-9274-y Vance GM, VanDyk JK, Rowley WA (1995) A device for sampling aquatic insects associated with carrion in water. J Forensic Sci 40:479–482 Voigt J (1965) Specific post-mortem changes produced by larder beetles. J Forensic Med 12:76–80 von Niezabitowski ER (1902) Experimentelle beiträge zur Lehre von der Leichenfauna [Experimental contributions to the science of the fauna of corpses]. Vierteljahrsschr gerichtl Med öffentl Sanitätsw 3:44–50 Walker TJ Jr (1957) Ecological studies of the arthropods associated with certain decaying materials in four habitats. Ecology 38:262–276 Wasti SS (1972) A study of the carrion of the common fowl, Gallus domesticus, in relation to arthropod succession. J Georgia Entomol Soc 7:221–229 Watson EJ, Carlton CE (2003) Spring succession of necrophilous insects on wildlife carcasses in Louisiana. J Med Entomol 40:338–347 Watson EJG (2004) Faunal succession of necrophagous insects associated with high-profile wildlife carcasses in Louisiana. PhD Thesis, Department of Entomology, Louisiana State University, Baton Rouge, LA, USA Wilson DS (1983) The effect of population structure on the evolution of mutualism: a field test involving burying beetles and their phoretic mites. Am Nat 121:851870 Wilson E (1844) Researches into the structure and development of a newly discovered parasitic animalcule of the human skin–the Entozoon folliculorum. Philos Trans R Soc Lond 134:305–319 Wolff M, Builes A, Zapata G et al (2004) Detection of Parathion (O, O-diethyl O-(4-nitrophenyl) phosphorothioate) by HPLC in insects of forensic importance in Medellı́n, Colombia. Anil Aggrawal’s Internet J Forensic Med Toxicol 5:6–11 Wyss C, Cherix D (2006) Traité d’entomologie forensique. Les insectes sur la scène de crime [Treatise on forensic entomology. The insects at the crime scene]. Presse polytechniques et universitaires romandes, Lausanne Yoder WA (1972) Acarina (Arthropoda: Arachnida) associated with selected Michigan Silphidae (Coleoptera). Michigan State University, East Lansing 123