Carcases and mites

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.

Full text

Carcases and mites
Henk R. Braig Æ M. Alejandra Perotti
Received: 5 June 2009 / Accepted: 16 June 2009 / Published online: 24 July 2009
! 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 Me
´
gnin’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
Exp Appl Acarol (2009) 49:45–84
DOI 10.1007/s10493-009-9287-6

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 Me
´
gnin 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 (Me
´
gnin 1894).
A short summary of the 8 waves was published a year later (Me
´
gnin 1895). There remains
an oddity in Me
´
gnin’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
Me
´
gnin 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
Me
´
gnin’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 decom-
position. 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.
Me
´
gnin’s appreciation of mites in a forensic context has been acknowledged early on by
forensic entomologists and pathologists (Graells 1886; Rı
´
os 1902a, b; Lecha-Marzo 1917;
Porta 1929). However, the proposed succession of insects and Me
´
gnin’s interpretations
were questioned over time by many (Strauch 1912; Wyss and Cherix 2006).
Me
´
gnin’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 Me
´
gnin’s species
descriptions and included them in their revisions.
Serrator amphibius Me
´
gnin (1894) is a revision by Me
´
gnin himself of Tyroglyphus
rostro-serratus Me
´
gnin 1873 and should now be recognised as
Histiostoma feroniarum
(Dufour 1839) (Histiostomatidae, Astigmata). The identification of Serrator necrophagus
Me
´
gnin (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 Me
´
gnin (1894) (? Uropodidae Kramer 1881,
Mesostigmata) and Trachynotus cadaverinus Me
´
gnin (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
46 Exp Appl Acarol (2009) 49:45–84
123

Table 1 Based on 15 years of experience at the Paris morgue, Me
´
gnin described 8 waves, squads or
periods of arthropod succession on human corpses exposed to the air (escouades or se
´
ries des travailleurs de
la mort [sections or series of death workers or gravediggers of nature (Gaudry 2002)])
Faunal succession as established by Me
´
gnin 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
Exp Appl Acarol (2009) 49:45–84 47
123

Table 1 continued
Faunal succession as established by Me
´
gnin 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 Me
´
gnin,=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
48 Exp Appl Acarol (2009) 49:45–84
123

Table 1 continued
Faunal succession as established by Me
´
gnin 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 Me
´
gnin) ?
Trachytidae
Uroseius acuminatus (=Trachynotus cadaverinus Me
´
gnin) ?
Astigmata
Acaridae
Acarus siro (=Tyroglyphus siro, Tyrolichus casei)
Tyrophagus longior (=Tyroglyphus longior, Tyroglyphus infestans)
Histiostomatidae
Histiostoma feroniarum (=Serrator amphibius Me
´
gnin, Tyroglyphus rostro-serratus
Me
´
gnin)
Serrator necrophagus Me
´
gnin ?
Glycyphagidae
Glycyphagus destructor (=Glyciphagus cursor Me
´
gnin, 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
Exp Appl Acarol (2009) 49:45–84 49
123

Me
´
gnin’s species and often not in easily accessible publications, which might have con-
tributed to them being overlooked (Rı
´
os 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, Leiod-
inychus 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 .
Me
´
gnin differentiates between Glyciphagus spinipes Ch. Rob. and Glyciphagus cursor
Me
´
gnin (1894), both are now considered synonyms of the pilous or groceries mite Gly-
cyphagus (Lepidoglyphus) destructor (Schrank 1781) (Glycyphagidae, Astigmata). Me
´
gnin
also differentiates between Tyroglyphus longior Gervais 1844 (Me
´
gnin 1894) and
Tyroglyphus infestans Berlese 1884 (Me
´
gnin 1898), both are now synonyms of the seed
mite Tyrophagus longior (Gervais 1844). However, the Tyrophagus species reported by
Me
´
gnin might have been a mixture of species (Perotti 2009).
The forensically important bulb mite species Cœpophagus echinopus depictured in
detail in Me
´
gnin’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) (Sams
ˇ
in
ˇ
a
´
k 1960). Tyroglyphus mycophagus Me
´
gnin
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 Me
´
gnin’s mite-rich sixth wave but have not been reported
since then (Lecha-Marzo 1917).
The carrion or grave fly, Ophyra cadaverina Me
´
gnin (1894) (Muscidae, Diptera), fifth
wave, had been ignored by entomologists for some time. Around 85 years after the original
publication in Me
´
gnin’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’.
Table 1 continued
Faunal succession as established by Me
´
gnin 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 Me
´
gnin (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
50 Exp Appl Acarol (2009) 49:45–84
123

The bottle also contained three specimens of O. cadaverina that allowed the identification
of Me
´
gnin’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 Schu
¨
hli et al. 2004, 2007). The bottle must have been part of original
material offered to the museum by Me
´
gnin. Acarologists have not yet investigated whether
some of the mites have been saved as well.
It is surprising that Me
´
gnin 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 pingui-
nalis, 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 envi-
ronmental 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 (Mi-
chelsen 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 Me
´
gnin, 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 [franc-
hement 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 desse
´
che
´
]’ but didn’t
assign specific species to each habitat (Leclercq and Verstraeten 1988a, 1993; Leclercq 2002).
The waves of arthropods in Me
´
gnin’s system overlap with each other; they often form a
continuum where it becomes difficult to say where one particular wave ends and a sub-
sequent 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 Me
´
gnin’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
Exp Appl Acarol (2009) 49:45–84 51
123

structured overview of the time line, the potential mite carriers and of the potential pre-
dators of mites still might be the use of Me
´
gnin’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
52 Exp Appl Acarol (2009) 49:45–84
123

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, Me
´
gnin 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, how-
ever, that the abundance was helped by special environmental circumstances’ (Me
´
gnin
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. Me
´
gnin’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
Exp Appl Acarol (2009) 49:45–84 53
123

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 (Me
´
gnin 1887,
1894). In the introduction to the section on the fauna of buried and entombed corpses,
Me
´
gnin 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; Pe
´
rez 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
54 Exp Appl Acarol (2009) 49:45–84
123

(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, Sph-
aeroceridae, 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 Sphae-
roseius 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
Exp Appl Acarol (2009) 49:45–84 55
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
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Fresh stage, initial decay
Mesostigmata
Arctoseius sp. Ascidae Pig On Grassland, bush Spain 12–2 Castillo Miralbes 2002
Haemogamasus sp. Haemogamasidae
a
Pig On Grassland, bush Spain 12–2 Castillo Miralbes 2002
Laelapidae Increasing Chicken Under Woods ME, USA Wasti 1972
Glyptholaspis
americana
Macrochelidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrocheles merdarius Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
M. muscaedomesticae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrochelidae Some Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998; Davis and Goff 2000
Increasing Chicken Under Woods ME, USA Wasti 1972
Parasitus sp. Parasitidae Pig On Grassland, bush Spain 12–2 Castillo Miralbes 2002
Poecilochirus
necrophori
High Mice On Forest MI, USA Wilson 1983
P. silphaphila Large carcases On MI, USA Yoder 1972; Brown and Wilson 1994
Parasitidae Increasing Chicken Under Woods ME, USA Wasti 1972
Fuscuropoda sp. Urodinychidae Abundant Chicken On Farm IA, USA Rives and Barnes 1988
Sp 1–3 Uropodidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Some Pig On Several HI, USA Hewadikaram and Goff 1991; Avila
and Goff 1998
Astigmata
Acarus farris Acaridae Dog Assoc. Costa Rica OConnor 2009
NY, USA OConnor 2009
A. siro Acaridae Common Lizard, chicken On Woods Nigeria Iloba and Fawole 2006
Acaridae Moderate Chicken Under Woods MA, USA Wasti 1972
Chicken On, under Field Spain 10–3 Arnaldos et al. 2004
56 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Oribatida
Oribatids Decline Chicken Under Woods MA, USA Wasti 1972
Prostigmata
Demodex brevis Demodecidae Decline Human On Normal fauna Worldwide Desch 2009
D. folliculorum Decline Human On Normal fauna Worldwide Desch 2009
Demodecidae Decline Most mammals On Normal fauna Worldwide Wilson 1844; Gmeiner 1908
Bdellidae Abundant Chicken Under Woods MA, USA Wasti 1972
Rhagidiidae Abundant Chicken Under Woods MA, USA Wasti 1972
Trombidiidae Moderate Chicken Under Woods MA, USA Wasti 1972
Pig On Bush Spain 12–2 Castillo Miralbes 2002
Putrefaction, bloated stage—terrestrial
Mesostigmata
Arctoseius sp. Ascidae Pig On Grassland, bush Spain 1–3 Castillo Miralbes 2002
Haemogamasus sp. Haemogamasidae Pig On Grassland, bush Spain 1–3 Castillo Miralbes 2002
Laelapidae Fewer Chicken Under Woods ME, USA Wasti 1972
Glyptholaspis
americana
Macrochelidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrocheles merdarius Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
M. muscaedomesticae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrochelidae Some Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998; Davis and Goff 2000
Macrochelidae Fewer Chicken Under Woods ME, USA Wasti 1972
Pachylaelaps sp. Pachylaelapidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Parasitus sp. Parasitidae Pig On Grassland, bush Spain 1–3 Castillo Miralbes 2002
Pergamasus sp. Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Some Pig On, under Several HI, USA Hewadikaram and Goff 1991
Parasitidae Fewer Chicken Under Woods ME, USA Wasti 1972
Common Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998
Exp Appl Acarol (2009) 49:45–84 57
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Poecilochirus sp. Some Rabbit On Woods CO, USA 7–8 De Jong and Chadwick 1999
Sp 1–3 Uropodidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Some Pig On Several HI, USA Hewadikaram and Goff 1991; Avila
and Goff 1998
Astigmata
Acarus siro Acaridae Common Fish, frog,
Lizard, chicken On Woods Nigeria Iloba and Fawole 2006
Lardoglyphus zacheri Lardoglyphidae Deer Assoc. UT, USA OConnor 2009
or Acaridae
Prostigmata
Trombidiidae Pig On Grassland, bush Spain 9,1–3 Castillo Miralbes 2002
Ixodida
Ixodidae Pig On Bush Spain 5 Castillo Miralbes 2002
Putrefaction, bloated stage—freshwater
Oribatida
Hydrozetes sp. Hydrozetidae Pig On Canada Hobischak and Anderson 2002
Black putrefaction, active decay
Mesostigmata
Cyrtolaelaps
mucronatus
Rhodacaridae One Human (26 d) On Forest Belgium 12 Leclercq 1978
Arctoseius sp. Ascidae Pig On Grassland, bush Spain 2–4 Castillo Miralbes 2002
Haemogamasus sp. Haemogamasidae Pig On Grassland, bush Spain 2–4 Castillo Miralbes 2002
Glyptholaspis
americana
Macrochelidae Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrocheles merdarius Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrochelidae Abundant Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998; Davis and Goff 2000
Macrocheles sp. 20, 2 Rat On, under Germany Scho
¨
nborn 1963
Pachylaelaps sp. Pachylaelapidae Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
58 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Parasitus stercorarius Parasitidae Common Pig, sparrow,
Taxonomic position
uncertain
Crow, mouse On Forest Germany Fourman 1936
Parasitus sp. Pig On Grassland, bush Spain 2–4 Castillo Miralbes 2002
Pergamasus sp.
b
Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Abundant Pig On, under Several HI, USA Hewadikaram and Goff 1991
Poecilochirus sp. Common Harbour seal On Rock MA, USA 5–10 Lord and Burger 1984a
Common Rabbit On Woods CO, USA 7–8 De Jong and Chadwick 1999
Parasitidae Abundant Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998
‘Gamasidae’ Large Guinea pig Under Woods W Australia 10–12 Bornemissza, 1957
Sp 1–3 Uropodidae Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Abundant Pig On Several HI, USA Hewadikaram and Goff 1991; Avila
and Goff 1998
Astigmata
Acarus siro Acaridae Common Fish, frog, pig
Sancassania berlesi Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Tyrophagus
putrescentiae
Abundant Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Abundant Pig On Several HI, USA Hewadikaram and Goff 1991
Acaridae Abundant Pig On Several HI, USA Avila and Goff 1998
Spinanoetus spp. nov Histiostomatidae Common crow,
White-tailed
deer
On MI, USA OConnor 2009
Pelzneria spp. nov Mouse, crow,
White–tailed
deer
On MI, USA OConnor 2009
‘Tyroglyphidae’ Some Guinea pig Under Woods W Australia 10–12 Bornemissza 1957
Exp Appl Acarol (2009) 49:45–84 59
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Oribatida
Galumna tarsipennata Galumnidae Chicken On, under Field Spain 2–3 Arnaldos et al. 2004
Zygoribatula connexa Oribatulidae Chicken On, under Field Spain 2–3 Arnaldos et al. 2004
Prostigmata
Trombidiidae Pig On Grassland, bush Spain 9–10,3–4 Castillo Miralbes 2002
Ixodida
Ixodidae Pig On Grassland, bush Spain 2–5 Castillo Miralbes 2002
Butyric fermentation, advanced decay
Mesostigmata
Arctoseius sp. Ascidae Pig On Grassland, bush Spain 10,4 Castillo Miralbes 2002
Asca sp. Scarce Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Proctolaelaps
epuraeae
Many Human (3 m) On, under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Proctolaelaps sp. ? Ten Human (2 m) On Belgium 12 Leclercq and Verstraeten 1988b
Zerconopsis remiger Many Human (3 m) Under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Hypoaspis aculeifer Laelapidae Many Human (3 m) On, under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Hypoaspis sp. ? Ten Human (2 m) On Belgium 12 Leclercq and Verstraeten 1988b
Glyptholaspis
americana
Macrochelidae Many Human (3 m) Under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrocheles glaber Fox On Garden England 10 Smith 1975
M. merdarius Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
M. muscaedomesticae One Human (17 d) On Small wood England 10 Easton and Smith 1970
Some Impala On Woods South
Africa
1–10 Braack 1986, 1987
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Macrocheles sp. Abundant Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Increase Chicken Under Woods ME, USA Wasti 1972
Pig On Woods SC, USA 8 Payne and Crossley 1966
60 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Macrochelidae Common Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998; Davis and Goff 2000
Gamasodes spiniger Parasitidae Fox On Garden England 10 Smith 1975
‘Gamasus’ sp. Many Human (2.7 y) On Cask Switzerland Hunziker 1919
Paragamasus sp. Many Human (3 m) Under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Parasitus fimetorum Fox On Garden England 10 Smith 1975
Parasitus sp. Abundant Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Increase Chicken Under Woods ME, USA Wasti 1972
Pig On Grassland Spain 4 Castillo Miralbes 2002
Pig On Woods SC, USA 8 Payne and Crossley 1966
Pergamasus sp. Scarce Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Phorytocarpais spp. Abundant Rabbit On, under Urban Alex.,
Egypt
11–4 Tantawi et al. 1996
Poecilochirus carabi Common Human (35 d) On Belgium 8 Leclercq and Verstraeten 1988b
Several Human (2 m) On beetle Pine forest Spain 11
Many Human
hanging
Under Salon
˜
a-Bordas pers. comm.
P. necrophori One Human (17 d) On Small wood England 10 Easton and Smith 1970
P. subterraneus Common Human (35 d) On Belgium 8 Leclercq and Verstraeten 1988b
Poecilochirus sp. Common Harbour seal On Rock MA, USA 5–10 Lord and Burger 1984a
Common Rabbit On Woods CO, USA 7–8 De Jong and Chadwick 1999
‘Gamasidae’ Large Guinea pig Under Woods W Australia 10–12 Bornemissza, 1957
Urobovella pulchella Uropodidae Many Human (3 m) On, under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Uroseius sp. Pig On Grassland, bush Spain 10 Castillo Miralbes 2002
Apionoseius sp. Discourellidae Medium Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Idendity unclear
Haemogamasus sp. Haemogamasidae Pig On Grassland, bush Spain 10,4 Castillo Miralbes 2002
Melittiphis ? sp. Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Exp Appl Acarol (2009) 49:45–84 61
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Gamasellus sp. Rhodacaridae or
Ologamasidae
Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Zercon sp. Zerconidae Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Astigmata
Myianoetus diadematus Histiostomatidae Mass occ. Human (1.3 y) On Basement Germany Russell et al. 2004
Acarus siro Acaridae Common Fish, frog, pig
Lizard, chicken On Woods Nigeria Iloba and Fawole 2006
Cosmoglyphus sp. Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Sancassania berlesei Many Human (3 m) Under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Sancassania sp. nov Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Sancassania sp. nov Deer, raccoon On USA OConnor 2009
Sancassania sp. Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Abundant Pig On Burial pit SC, USA 6–11 Payne et al. 1968
Tyrophagus
putrescentiae
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Common Pig On Several HI, USA Hewadikaram and Goff 1991
Lardoglyphus zacheri Lardoglyphidae or
Acaridae
Bird Feathers
under
TX, USA OConnor 2009
‘Tyroglyphidae’ Immense Guinea pig Under, on Woods W Australia 10–12 Bornemissza 1957
Acaridae Common Pig On Several HI, USA Avila and Goff 1998
Oribatida
Platynothrus peltifer Camisiidae Many Human (3 m) Under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Minunthozetes
semirufus
Mycobatidae Many Human (3 m) Under Deciduous forest Spain 8 Salon
˜
a et al. in prep.
Malacanothrus sp. Malacanothridae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Ceratoppia bipilis Ceratoppiidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Liacaridae Common Pig On Several HI, USA Hewadikaram and Goff 1991
Medioppia pinsapi Oppiidae Chicken On, under Field Spain 10–12 Arnaldos et al. 2004
62 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Oribatula tibialis Oribatulidae Chicken On, under Field Spain 10–12 Arnaldos et al. 2004
Oribatida spp. Common Pig On Several HI, USA Davis and Goff 2000
Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Prostigmata
Lephus spp. Erythraeidae Pig On Woods SC, USA 8 Payne and Crossley 1966
Penthaleus major Eupodidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Trombidiidae Pig On Grassland, bush Spain 10 Castillo Miralbes 2002
Ixodida
Dermacentor variabilis Ixodidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Ixodidae Pig On Bush Spain 4 Castillo Miralbes 2002
Dry decomposition, skeletal stage
Mesostigmata
Leiodinychus krameri Dinychidae
(Uropodidae)
Myriads Human ([1 y) On, in Cellar France Me
´
gnin 1894
Common Human On Canada Johnston and Villeneuve 1897
Holostaspis sp. Laelapidae Common Human (11 y) On Grave DC, USA Motter 1898
Hypoaspis sp. Common Human
(20–30 y)
On Grave DC, USA Motter 1898
Common Dog (3 m) On Grave DC, USA Motter 1898
Laelaps (Iphis) sp. Common Human (11 y) On Grave DC, USA Motter 1898
Melittiphis ? sp. Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Glyptholaspis
americana
Macrochelidae Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA Early and Goff 1986; Goff 1989
Macrocheles glaber One Human (3 m) On Belgium 6 Leclercq and Verstraeten 1988b
M. merdarius Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Exp Appl Acarol (2009) 49:45–84 63
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
M. muscaedomesticae Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Some Impala On Woods Sth Africa 1–10 Braack 1986, 1987
Macrocheles sp. Abundant Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Abundant Small animals
f
On Oak forest IL, USA 4–11 Johnson 1975
Macrochelidae Common Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998; Davis and Goff 2000
Pachylaelaps sp. Pachylaelapidae Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
‘Gamasus’ sp. Parasitidae Common Human
(30–40 y)
On Grave DC, USA Motter 1898
Common Dog (3 m) On Grave DC, USA Motter 1898
Parasitus sp. Abundant Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Common Small animals
f
On Oak forest IL, USA 4–11 Johnson 1975
Pergamasus sp.
c
Scarce Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Abundant Small animals
f
On Oak forest IL, USA 4–11 Johnson 1975
Poecilochirus sp. Common Rabbit On Woods CO, USA 7–8 De Jong and Chadwick 1999
Uroseius acuminatus Trachytidae Common Human (3 y) On, in France 5 Me
´
gnin 1894
Uropoda depressa Uropodidae Common Human (3–7 y) On Grave DC, USA Motter 1898
Identity unclear
Uropoda sp. Common Dog (3–5 m) On Grave DC, USA Motter 1898
Sp 1–3 Uropodidae Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Common Pig On Several HI, USA Hewadikaram and Goff 1991; Avila and
Goff 1998
Asca sp. Ascaidae Scarce Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Apionoseius sp. Discourellidae Medium Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
Identity unclear
Gamasellus sp. Rhodacaridae Rare Dog In, on, under Woods, pasture TN, USA 1–12 Reed 1958
or Ologamasidae
64 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
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
Astigmata
Acarus immobilis Acaridae Common Human (1.3 y) On Basement Germany Russell et al. 2004
Raccoon On OH, USA OConnor 2009
A. siro Common Human (3 y) On, in Rural France 5 Me
´
gnin 1894
Myriads Human ([1 y) On, in Cellar France Me
´
gnin 1894
Common Fish, frog, pig
Lizard, chicken On Woods Nigeria Iloba and Fawole 2006
Acarus (Tyroglyphus)
sp.
Common Human (3 y) On Grave DC, USA Motter 1898
Cosmoglyphus sp. Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Rhizoglyphus
echinopus
Myriads Human ([1 y) On, in Cellar France Me
´
gnin 1894
Abundant Human (2 y) On Urban France 10 Me
´
gnin 1894
Common Human (2–3 y) Bulbs of lily Garden, burrial France Me
´
gnin 1894
Sancassania berlesei Abundant Human
(3–8 m)
On, in Urban France 1 Brouardel 1879
Abundant Human (7–8 y) On House France Me
´
gnin 1894
784 Human (3 m) On Belgium 6 Leclercq and Verstraeten 1988b
Abundant Human (3.5 m) On Belgium 1 Leclercq and Verstraeten 1988b
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Sancassania sp. nov Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Sancassania sp. nov Deer, raccoon On USA OConnor 2009
Sancassania sp. Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Tyrophagus longior Abundant Human
(3–8 m)
On, in Urban France 1 Brouardel 1879
Abundant Human (7–8 y) On House France Me
´
gnin 1894
Very rare Human (1 y) On House France 1 Me
´
gnin 1894
Exp Appl Acarol (2009) 49:45–84 65
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Common Human (3 y) On, in Rural France 5 Me
´
gnin 1894
Myriads Human ([1 y) On, in Cellar France Me
´
gnin 1894
Myriads Human On House, trunk France Me
´
gnin 1898
Common Human On, in Canada Johnston and Villeneuve 1897
T. putrescentiae Abundant Human (1.3 y) On Basement Germany Russell et al. 2004
Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Common Pig On Several HI, USA Hewadikaram and Goff 1991
Tyrophagus sp. Common Human (18 m) On, around House France 3 Me
´
gnin 1894
Common Human (2 y) On Urban France 6 Me
´
gnin 1894
T. (Hypopus) sp. Common Human
(20–71 y)
On Grave DC, USA Motter 1898
Identity unclear
Acaridae Few Human
(summer)
Alps France 10 Leclercq and Verstraeten 1992
Common Pig On Several HI, USA Avila and Goff 1998
Glycyphagus destructor Glycyphagidae Very rare Human (1 y) On House France 1 Me
´
gnin 1894
Common Human On France Me
´
gnin 1894
Glycyphagidae Large Human (28 y) On Embalmed MI, USA Merritt et al. 2007
Histiostoma feroniarum Histostomatidae Common Human On France Me
´
gnin 1894
Common Human On Canada Johnston and Villeneuve 1897
H. necrophagus Common Human On France Me
´
gnin 1894
Composite species,
unrecognisable
Common Human On Canada Johnston and Villeneuve 1897
H. sachsi Two Human (3 m) On Belgium 6 Leclercq and Verstraeten 1988b
Histiostoma sp. One Human (3 m) On Belgium 6 Leclercq and Verstraeten 1988b
Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
66 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Myianoetus ? sp. Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Histostomatidae Common Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998
Lardoglyphus
radovskyi
Lardoglyphidae Human Pelvis, Mummy NV, USA Baker 1990
or Acaridae Gut content Radovsky 1970
L. robustisetosus Human Gut content Mummy Chile Baker 1990
L. zacheri Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Racoon On USA OConnor 2009
Czenspinskia
transversostriata
Winterschmidtiidae Common Human (53 d) Soil Clothing HI, USA 3–5 Goff 1991
Common Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Oribatida
Aphelacarus acarinus Aphelacaridae Human Remains Tomb Spain Hidalgo-Argu
¨
ello et al. 2003
Hoplophora (Tritia) sp. Euphthiracaridae Common Human On Grave DC, USA Motter 1898
Platynothrus peltifer Camisiidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Ceratoppia bipilis Ceratoppiidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Galumnidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Rostrozetes spp.
d
Haplozetidae Common Skin of animals On
Haplozetidae 238 Rat On Campus Cameroon 2–3 Feugang Youmessi et al. 2008
Liacaridae Common Pig On Several HI, USA Hewadikaram and Goff 1991
Malacanothrus sp. Malacanothridae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Oribatida spp. Common Pig On Several HI, USA Davis and Goff 2000
Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Increase Chicken Under Woods MA, USA Wasti 1972
Prostigmata
Cheyletus eruditus Cheyletidae Abundant Human ([1 y) On Cellar France Me
´
gnin 1894
Exp Appl Acarol (2009) 49:45–84 67
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Podapolipidae Human Remains Tomb Spain Hidalgo-Argu
¨
ello et al. 2003
Tarsonemoidea Human Remains Tomb Spain Hidalgo-Argu
¨
ello et al. 2003
Tarsotomus sp. nov Anystidae Abundant Rabbit On, under Urban Alexandria,
Egypt
7–8 Tantawi et al. 1996
Erythraeus sp. Erythraeidae Pig On Woods SC, USA 8 Payne and Crossley 1966
Penthaleus major Eupodidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Trombidium sp. Trombidiidae Common Small animals
f
On Oak forest IL, USA 4–11 Johnson 1975
Ixodida
Dermacentor variabilis Ixodidae Few Dog Under Woods, pasture TN, USA 1–12 Reed 1958
Pig On Woods SC, USA 8 Payne and Crossley 1966
Undetermined stage
Mesostigmata
Epicrius mollis Epicriidae Male Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
E. thanathophilus General,
human?
On Porta 1929
Celaenopsis cuspidatus Celaenopsidae General,
human?
On Porta 1929
Cornigamasus lunaris Parasitidae Few Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Holoparasitus
calcaratus
Few Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Paracarpais furcatus General,
human?
On Porta 1929
Pergamasus crassipes Few Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Pergamasus sp.
e
Hundreds Rat On Copse, grassland England 8–12 Collins 1970
Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Poecilochirus sp. Common Rat On Field CO, USA 7–8 De Jong and Hoback 2006
Parasitidae Common Bear, deer,
Alligator, pig On LA, USA Watson 2004
68 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Gamasida Some Pig On Several HI, USA 1–4 Davis and Goff 2000
Asca craneta Ascidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Gamasellodes bicolor Few Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Iphidozercon gibbus Some Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Proctolaelaps sp. nov Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Zerconopsis
decemremiger
Some Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Ascidae Some Pig On, under Several HI, USA Richards and Goff 1997; Avila and Goff
1998
Digamasellidae Some Pig On Several HI, USA Avila and Goff 1998
Digamasellidae As control Turtle Under Woods MA, USA 6–8 Abell et al. 1982
Diplogyniidae Increasing Turtle Under Woods MA, USA 6–8 Abell et al. 1982
Eviphidae Some Pig On Several HI, USA Avila and Goff 1998
Hypoaspis
(Cosmolaelaps)
vacua
Laelapidae Some Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Laelapidae 146, large Turtle On, under Woods MA, USA 6–8 Abell et al. 1982
Increase Rat Under Copse, grassland England 8–12 Collins 1970
Some Pig Associated Several HI, USA Richards and Goff 1997
Macrocheles sp. Macrochelidae Hundreds Rat On Copse, grassland England 8–12 Collins 1970
Macrocheles sp. N Rodents On USA Krantz and Whitaker 1988
M. agilis Carrion In Australia Halliday 2000
M. lagodekhensis group Several Roe deer On Slovakia 4, 5,7 Mas
ˇ
a
´
n 1993
M. matrius Weasel On MT, USA Krantz and Whitaker 1988
M. mykytowyczi Fish, squid,
carrion
In Forest Australia Halliday 2000
M. nataliae (=melisii) Vole On Lithuania Hyatt and Emberson 1988
Small
mammals
On USSR Bregetova and Koroleva 1960
Exp Appl Acarol (2009) 49:45–84 69
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
M. peckorum Carrion trap On Australia Halliday 2000
M. spatei Squid On Australia Halliday 2000
M. tessellatus Squid On Australia Halliday 2000
Macrochelidae Increase Rat Under Copse, grassland England 8–12 Collins 1970
Ologamasidae Some Pig Associated Several HI, USA Richards and Goff 1997
Paraholaspidae Some Pig Associated Several HI, USA Richards and Goff 1997
Phytoseius hawaiiensis Phytoseiidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Phytoseiidae Increasing Turtle Far, under Woods MA, USA 6–8 Abell et al. 1982
Podocinidae Some Pig Associated Several HI, USA Richards and Goff 1997
Rhodacaridae Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Trachytes aegrota Trachytidae Several Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Uropodidae As control Turtle Under Woods MA, USA 6–8 Abell et al. 1982
Increase Rat Under Copse, grassland England 8–12 Collins 1970
Veigaia nemorensis Veigaiidae Few Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Veigaiidae Some Pig On Several HI, USA Richards and Goff 1997; Davis and Goff
2000
Prozercon kochi Zerconidae Few Small animal On Alder forest Poland 8 Gwiazdowicz and Klemt 2004
Astigmata
Acaridae As control Turtle Under Woods MA, USA 6–8 Abell et al. 1982
Lardoglyphus sp. Lardoglyphidae or
Acaridae
Some Impala On Woods South
Africa
1–10 Braack 1986
Astigmata Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Oribatida
Achipteriidae Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Ceratozetidae Increasing Turtle Under Woods MA, USA 6–8 Abell et al. 1982
Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Eremaeidae One Rat On Field CO, USA 7–8 De Jong and Hoback 2006
Euzetidae Decrease Rat Under Copse, grassland England 8–12 Collins 1970
70 Exp Appl Acarol (2009) 49:45–84
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Galumnidae Increasing Turtle Far, under Woods MA, USA 6–8 Abell et al. 1982
Hypochthoniidae Increasing Turtle Far, under Woods MA, USA 6–8 Abell et al. 1982
Mycobatidae Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Nothridae 3, small Turtle On, under Woods MA, USA 6–8 Abell et al. 1982
Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Oppiidae Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Oribatulidae Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Prostigmata
Erythraeus sabulosus Erythraeidae General,
human?
On Porta 1929
Anystidae Few Rat On Field CO, USA 7–8 De Jong and Hoback 2006
Bdellidae Some Pig Associated Several HI, USA Richards and Goff 1997
Camerobiidae Some Pig On Several HI, USA Avila and Goff 1998
Cunaxa sp. Cunaxidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Ereynetidae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Some Pig On Several HI, USA Avila and Goff 1998
Eupodidae Some Pig Associated Several HI, USA Richards and Goff 1997
Pygmephorus sp. Pygmephoridae Some Impala On Woods South
Africa
1–10 Braack 1986
Pygmephoridae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Some Pig On Several HI, USA Avila and Goff 1998
Rhagidiidae Increasing Turtle Under Woods MA, USA 6–8 Abell et al. 1982
Some Pig Associated Several HI, USA Richards and Goff 1997
Scutacaridae Some Cat On, under Xero ? mesophytic HI, USA 3–5 Early and Goff 1986; Goff 1989
Tarsonemoidea Some Pig On Several HI, USA Avila and Goff 1998
Prostigmata Decrease Rat Under Copse, grassland England 8–12 Collins 1970
Endeostigmata
Terpnacaridae Some Pig On Several HI, USA Avila and Goff 1998
Exp Appl Acarol (2009) 49:45–84 71
123

Table 3 continued
Species Family Abundance Host Location Habitat Country Season
(month)
Reference
Ixodida
Dermacentor variabilis Ixodidae Few Bear On LA, USA 12–2 Watson 2004
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)
72 Exp Appl Acarol (2009) 49:45–84
123

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 cate-
gory. 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; Oeco-
phoridae; 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
Exp Appl Acarol (2009) 49:45–84 73
123

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 decom-
position characterised by the astigmatid mites Tyrophagus putrescentiae and Acarus im-
mobilis; 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 decomposi-
tion. 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 dis-
covery 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 rep-
resentative 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.
74 Exp Appl Acarol (2009) 49:45–84
123

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 dra-
matic 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).
Exp Appl Acarol (2009) 49:45–84 75
123

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 heli-
ophilic 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). Mac-
rocheles 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 asso-
ciations 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, ele-
phants, rhinoceroses, mole rats, whales and hippopotamuses share naturally the nakedness
76 Exp Appl Acarol (2009) 49:45–84
123

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 ori-
batid 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 Ku
¨
hnelt (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 geo-
graphical 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 (Macroc-
helidae, 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 cm
3
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
Exp Appl Acarol (2009) 49:45–84 77
123

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; Mendonc¸a 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 acaro-
logical 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. diadem-
atus, 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 Lever-
hulme Trust. Additional information was kindly provided by M. Lee Goff, Paola Magni, Marta I. Salon
˜
a-
Bordas and Francis D. Feugang Youmessi. The authors like to thank Marilo
´
Moraza and Barry M. OConnor
for advice and reviewing an earlier version of the manuscript.
78 Exp Appl Acarol (2009) 49:45–84
123

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 sarcosappro
´
faga de la Regio
´
n de Murcia. Su aplicacio
´
na
la mediciona legal [Studies on the sarcosaphrophagous fauna in the Region of Murcia; its application in
legal medicine]. Departamento de Biologı
´
a Animal. Universidad de Murcia, Murcia
Athias-Binche F (1994) La Phore
´
sie chez les Acariens—Aspects Adaptatifs et Evolutifs [Phoresy in aca-
rina—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. De
´
couverte du cadavre d’un enfant
nouveau-ne
´
dans une chemine
´
e ou
`
il s’e
´
tait momifie
´
. De
´
termination de l’e
´
poque de la naissance par la
pre
´
sence de nymphes et de larves d’insectes dans le cadavre et par l’e
´
tude de leurs me
´
tamorphoses
[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
Me
´
d Le
´
g 4(se
´
rie):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: Para-
sitidae) 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, He
´
douin 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 de
´
termination de l’e
´
poque de la naissance et de la mort d’un nouveau-ne
´
e, faite a
`
l’aide de la pre
´
sence des acares et des chenilles d’aglosses dans cadavre momifie
´
[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 Me
´
d Le
´
g 2(se
´
rie 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 cada
´
veres en el Alto Arago
´
n (Espan
˜
a)
[Study of the entomofauna associated with corpses in the region of Alto Arago
´
n (Spain)]. Sociedad
Entomolo
´
gica Aragonesa, Zaragoza
Exp Appl Acarol (2009) 49:45–84 79
123

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 eco-
system. 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 u
¨
ber die Lebensweise wirbelloser Aasfresser [Comperative
studies on the ecology of invertebrate carrion feeders]. Sitzungsb Ko
¨
nigl 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, Djie
´
to-Lordon C, Gaudry E et al (2008) Contribution to the research of the ento-
mological 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. Me
´
gnin 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: Dusba
´
bek 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
80 Exp Appl Acarol (2009) 49:45–84
123

Goff ML, Garcı
´
a Garcı
´
a MD, Arnaldos Sanabria MI (2004) Entomologı
´
a cadave
´
rica: Fundamentos y
aplicacio
´
n. Referencia a la entomologı
´
a espan
˜
ola [Forensic entomology: basics and applications. A
reference to Spanish entomology]. In: Gisbert Calabuig JA, Villanueva Can
˜
adas E et al (eds) Tratado
de Medicina Legal y Toxicologı
´
a [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 Fı
´
s 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-Argu
¨
ello MR, Dı
´
ez Ban
˜
os N, Fregeneda Grandes J et al (2003) Parasitological analysis of Leonese
royalty from Collegiate-Basilica of St. Isidoro, Le
´
on (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 sarcosapro
´
faga en
Co
´
rdoba (Argentina): aplicacio
´
n a la entomologı
´
a forense [Preliminary study of the sarcosaprophytic
community in Cordoba (Argentina): applied to forensic entomology]. An Biol 27:191–201
Hunziker H (1919) U
¨
ber die Befunde bei Leichenausgrabungen auf den Kirchho
¨
fen Basels. Unter beson-
derer Beru
¨
cksichtung der Fauna und Flora der Gra
¨
ber [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, Dermanys-
soidea). Am Mus Novit 3151:1–8
Ku
¨
hnelt 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 Clı
´
nica, Madrid, pp 79–90
Leclercq M (1978) Entomologie et Me
´
decine Le
´
gale: Datation de la Mort [Entomology and forensic
medicine: dating the time of death]. Masson, Paris
Leclercq M (2002) L’entomologie le
´
gale 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 me
´
dicine le
´
gale. Datation de la mort: insectes et autres
arthropodes trouve
´
s sur les cadavres humains [Entomology and forensic medicine, determination of the
Exp Appl Acarol (2009) 49:45–84 81
123

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 me
´
decine le
´
gale. Datation de la mort. Acariens trouve
´
s
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 be
´
ne
´
voles dans les e
´
cosyste
`
mes terrestres:
observation ine
´
dite [Voluntary entomological street sweepers in the terrestrial ecosystems: a new
observation]. Notes Faun Gembloux 25:17–22
Leclercq M, Verstraeten C (1993) Entomologie et me
´
decine le
´
gale. L’entomofaune des cadavres humains:
sa succession par son interpre
´
tation, ses re
´
sultats, 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, Mendonc¸a 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 a
`
la me
´
decine
le
´
gale [A new case of the application of entomology in legal medicine]. Ann Hyg Publ Me
´
d Le
´
g
13(se
´
rie 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 black-
backed 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
Mas
ˇ
a
´
n 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 entomo-
toxicological examinations to identify causes of death EAFE Meeting 2008, Kolymbari, Greece
Me
´
gnin P (1887) La faune des tombeaux [The fauna of graves]. C R Hebd Acad Sci 105:948–951
Me
´
gnin P (1894) La Faune des Cadavres. Application de l’Entomologie a
`
la Me
´
decine Le
´
gale [The fauna of
corpses. Application of entomology to forensic medicine]. G. Masson and Gauthier-Villars et Fils,
Paris
Me
´
gnin P (1895) La faune des cadavres [The fauna of carcasses]. Ann Hyg Publ Me
´
d Le
´
g se
´
rie 3(33):64–67
Me
´
gnin 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
Mendonc¸a de Souza SMF, Reinhard KJ, Lessa A (2008) Cranial deformation as the cause of death for a
child from the Chillon River Valley, Peru. Chungara
´
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
Pe
´
rez SP, Duque P, Wolff M (2005) Successional behavior and occurrence matrix of carrion-associated
arthropods in the urban area of Medellı
´
n, Colombia. J Forensic Sci 50:448–454
82 Exp Appl Acarol (2009) 49:45–84
123

Perotti MA (2009) Me
´
gnin 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) De
´
couverte de quelques insectes de J.-P. Me
´
gnin; identite
´
d’Ophyra cadaverina
Me
´
gnin (1894) (Diptera, Muscidae) [Discovery of several insects of J.-P. Me
´
gnin; identity of Ophyra
cadaverina Me
´
gnin (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
Rı
´
os T (1902a) Los insectos y la putrefaccio
´
n de los cada
´
veres [Insects and the decomposition of corpses]
(I-II). Clı
´
n Mod Rev Med Cirug 1:74–80
Rı
´
os T (1902b) Los insectos y la putrefaccio
´
n de los cada
´
veres [Insects and the decomposition of corpses]
(III-VI). Clı
´
n 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 Go
¨
rlitz 76:51–56
Sams
ˇ
in
ˇ
a
´
k K (1960) U
¨
ber einige myrmekophile Milben aus der Familie Acaridae [On some myrmecophylic
mites in the family Acaridae]. C
ˇ
as C
ˇ
esk Spol Entomol 57:185–192
Schnell e Schu
¨
hli 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 Schu
¨
hli 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
Scho
¨
nborn W (1963) Vergleichende zoozo
¨
notische 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
Exp Appl Acarol (2009) 49:45–84 83
123

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 o
¨
ffentl Sanita
¨
tsw
43:44–49
Strauch C (1928) Beitra
¨
ge zur natu
¨
rlichen 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 beitra
¨
ge zur Lehre von der Leichenfauna [Experimental
contributions to the science of the fauna of corpses]. Vierteljahrsschr gerichtl Med o
¨
ffentl Sanita
¨
tsw
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 car-
casses 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) phosp-
horothioate) by HPLC in insects of forensic importance in Medellı
´
n, Colombia. Anil Aggrawal’s
Internet J Forensic Med Toxicol 5:6–11
Wyss C, Cherix D (2006) Traite
´
d’entomologie forensique. Les insectes sur la sce
`
ne 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 (Cole-
optera). Michigan State University, East Lansing
84 Exp Appl Acarol (2009) 49:45–84
123