Fig. 1. A selection of fungal spores found in palynological preparations in a rape case investigated for Wiltshire Constabulary in 2009 [41,42]: (A) Clasterosporium exum; (B)
Diplocladiella scalarioides; (C) Camposporium cambrense; and (D) Dictyosporium toruloides. Not to scale. Photographs by J.A. Webb.
D.L. Hawksworth, P.E.J. Wiltshire / Forensic Science International 206 (2011) 111 4
on the surface of human cadavers. One key reference text for
forensic pathologists reports extensive growths on a boy after 6
weeks [60], while another refers to development after several
months on an embalmedcadaver [61]. However, the fungi involved
have not generally been identied even to genus, nor considered as
a forensic tool.
The rst researchers to appreciate that fungal growth on
corpses had a role in the determination of time of death were van
de Voorde and van Dijck [62]. These authors made isolations from
fungal growths on an eyelid and on inguinal skin of a murdered
baroness found in a room in Belgium. They incubated the fungal
isolates at the same temperature as that in which the body had
been found (a constant 12 8C controlled by a thermostat in the
room), and measured the colony sizes daily. They estimated that
the woman had died at least 18 days before her body was
discovered, and that agreed exactly with a subsequent admission
by the murderer. They considered that supercial fungi could help
in determining the time of death, when this was 1020 days
earlier, provided those temperature data were available. The fungi
involved in this case were named as: Cladosporium sp., Fusarium
sp., Geotrichum candidum, Hormodendron sp., Mortierella sp., and
Penicillium chysogenum (as P. notatum).
Unaware of the Belgian case, Ishii et al. [63] reported on fungal
growths found on the surface of a mummied body found in an
abandoned house, and also on skeletal remains discovered in a
forest, that they suggested may reveal local habitats. They
identied these as Aspergillus chevalieri (as Eurotium chevalieri), A.
repens (syn. E. repens) that was the most abundant, A. rubrum (as E.
rubrum), and Gliocladium sp. (probably actually a species of
Clonostachys). These authors recognised that more cases were
needed to establish the approach as a forensic tool, but did not
attempt to estimate PMI in these cases. They subsequently
reported on white growths dotted on the face of a male corpse
found in a kneeling position and partly in water at the bottomof an
open well in Japan [64]. The humidity was nearly 100%, and the
temperature constant at 1213 8C; the fungi were isolated into
culture and identied as Aspergillus terreus and Penicillium sp. The
deceased had been dead for 10 days, and although no growth
measurements of the colonies were made, they suggested that the
fungal data were consistent with the time as the fungi generally
colonise [in] 37 days a generalization based on the time-lag in
attacking foodstuffs [65]. These workers independently suggested
that fungi could provide a useful means of estimating PMI where
forensic entomological data were not available.
Menezes et al. [66] considered that Ishii et al.s claim of fungal
growths being of value in estimating PMIs would have been more
convincing if they had investigated the growths empirically. Their
response reafrmed their view of the potential, acknowledging
that there were unsolved problems regarding growth rates and the
actual fungi [67]. In a rejoinder, Menezes et al. [68] stressed the
need for more forensic cases to be documented from different
climatic conditions, and for experimental work on a substitute
subject, especially in relation to the pattern and rate of growth of
fungal species on cadavers.
When the body of a man was found by a tube line in Ruislip,
north-west London, the medical examiner considered it had been
there not more than 48 h. There was no evidence of any scavenger
or insect activity. However, botanical evidence, and a large circular
fungal colony under the chin, indicated that the body had lain in
situ for between 3 and 5 weeks [69]. It later transpired that the
actual time was 4 weeks 3 days. Decomposition had been delayed
because the weather had been exceptionally cold, and the low
temperatures had inhibited colonisation by ies. Security fencing
had excluded scavenging animals and the body had remained
intact.
In the case of the serial killing of prostitutes around Ipswich,
Suffolk at the end of 2006, two of the bodies were recovered from
water. These were ensheathed in fungal growth that had trapped
silt particles that had become graded as they accumulated in the
hyphal
10
weft. Several fungi were involved, including species of
Fusarium, Geotrichum, Mucor, and Pythium. The rst body that was
retrieved was completely ensheathed. With the second body, the
sheath was present though it was patchy and had collapsed due to
the much greater length of the hyphae than on the other body.
Where there had been skin slippage and exposure of the
underlying dermis, there had been fresh fungal colonisation on
some of the exposed areas. The fungi were intermixed and not
identied to species. Even though information on the growth of
these fungi in cold water was not available, it was obvious that the
body of this victim had been in the water longer than that of the
other. An estimate of at least 5 weeks submersion was suggested
by the condition of the fungal mycelium and other features of this
corpse. Using the same criteria, it was suggested that the woman
who had been found rst had been in the water for about 2 weeks.
These estimates agreed with the length of time the victims had
been reported to be missing [70,71].
Fungal colonies on, or associated with, human cadavers can give
indications of time since death as there is information on growth
rates of many moulds. But the reliability of any estimates will
depend on the accuracy of the identication of the fungus, the
storage methods for the body, and the availability of data on the
temperature and humidity at the site. As yet there are few precise
data on actual rates of growth on dead human tissues, especially
under different conditions of temperature and humidity. This
means that it is necessary to undertake experiments mirroring, as
near as is possible, the environmental parameters with which the
corpse was associated, using the actual strains found. We are
currently involved in such experimental work.
In a case undertaken for Tayside Police in 2009 [72], a man had
been repeatedly stabbed and died in a closed at so that ies were
excluded. Fungal colonies had developed on parts of the carpet and
on a sofa that had become soaked with body uids (Fig. 3). After the
discovery and removal of the body, data loggers were placed in the
room and consistently gave relative humidity readings of 3034%.
Colonies were measured in situ and subcultures made onto
articial media. Samples of the carpet were then removed and
kept dry for observation; no subsequent growth occurred within 5
days. This result was expected since most mould fungi require at
least 95% relative humidity for growth [73]. When the carpet
samples were re-wetted with bovine blood and kept at a high
humidity, there was a spurt of newfungal growth. A comparison of
the sizes of the new colonies, both in culture and on the carpet,
with those at the crime scene, suggested that the death had
occurred about 5 days prior to discovery of the body; this was
consistent with a subsequent admission of guilt. The three
Fig. 3. Carpet saturated with body uids and then dried, showing fungal colonies.
The large, dark greyish colonies are of Mucor plumbeus. See text for details.
Fig. 5. Xanthoria parietina growing on Sambucus nigra (elder) twigs: (A) natural
growth in full illumination; (b) natural growth in deep shade (underside of twig);
(C) sample grown in full illumination but then covered with a weighted object
wrapped in opaque plastic for 5 days. Not to scale.
D.L. Hawksworth, P.E.J. Wiltshire / Forensic Science International 206 (2011) 111 6
16th October in the previous year [79]. Similarly, as ergots
(Claviceps purpurea) developing on cereals and other grasses in the
autumn have hooks at the tips that facilitate attachment to
clothing or animal coats, their discovery would also suggest an
incident in the autumn.
Many mushrooms are also seasonal, and this is not only a
matter of species that form sporophores in the autumn. Some are
much more restricted, for example, the scarlet-cupped Sarcoscypha
species appear in the early spring on fallen twigs. These fungi have
distinctive spores and might provide important trace evidence if
trodden on by suspects. Another potentially useful fungus is the
mushroom Flammulina velutipes, that only appears after the rst
frosts of winter.
Fungal colonies grow in a circular manner when on solid
substrates, and the growth rates on articial media are often cited
in species descriptions. Those with non-septate hyphae belonging
to the Mucorales (pin-moulds) are, in general, much more rapidly
growing than those belonging to other groups (e.g. ascomycetes,
basidiomycetes) that have repeatedly septate hyphae. In a recent
case for Hertfordshire Constabulary, a mutilated murder victim
was deposited on his ventral surface on a steep bank above a small,
sluggish stream. The absence of blood indicated that the murder
and mutilation had occurred elsewhere. Small colonies of non-
sporing Mucor hiemalis had developed on the abdomen. Palyno-
logical analysis showed that mud that covered the corpse had been
derived from the stream and the victim had obviously been
dropped in or dragged through it. The colony size and immaturity
of the fungus indicated 12 days growth, and the pristine condition
of the myceliumshowed that it had developed in situ after the body
had been dragged through the stream. Experience has shown that
fungi, such as Mucor species, are unable to colonise and grow on
freshly dead skin and it would appear, on average, that this is
unable to happen for at least a week after death. This reinforced the
hypothesis that the victim had been killed and stored elsewhere
and had lain at the deposition site for a short time [80,81].
6. Cause of death, hallucinations, or poisoning
Mushroom poisoning can be accidental or deliberate, and may
be fatal. In most cases this arises fromthe consumption of wrongly
identied mushrooms by untrained collectors who often belong to
the family of the victim(s). In most cases the results are not fatal
but vary according to the amount consumed and the tolerance of
the individuals. In some cases the onset of symptoms is rapid,
while in others, involving kidney damage, they may not be evident
for several days. Most toxic are the fungi producing amanitins (e.g.
Amanita virosa), gyromitrin (Gyromitra esculenta), muscarine (e.g.
certain species of Conocybe and Inocybe), and orellanine (Cortinar-
ius orellanus), where only small amounts can prove fatal [82]. The
number of truly poisonous mushroom species is small, although
more can cause gastric problems in some individuals. Unfortu-
nately some of the poisonous species are rather common and, to
the non-specialist, can appear to be similar to certain edible
species. In the case of any suspected mushroom poisonings, any
remaining mushrooms should be secured, and stomach contents
analysed.
There are numerous books on mushroom identication, but
regional guides are seldom comprehensive as there are just so
many species. Consequently, of particular value in the case of
suspected poisonings are those dealing exclusively with the
identication of poisonous species [83,84], and a comprehensive
list of species known to have particular adverse effects has been
compiled [85]. When no intact mushrooms are available for
examination, spores and other microscopic fungal remains in
stomach and gut contents can be used to determine the species
consumed. A key for the identication of fragments of selected
hallucinogenic species has been compiled [86], and an on-line
identication program constructed [87,88]. Species-level identi-
cationis necessary to ensure the correct diagnosis and also that any
prescribed treatment is appropriate. In some continental European
countries where mushrooms are regularly collected and eaten,
poison control centres have accesses to mycological specialists or
have local specialist controllers (e.g. Switzerland).
The use of fungi as hallucinogens, neurotropic or psychoactive
drugs, has its roots in antiquity in both the Old and the New World
[89,90]. In all countries that have signed the 1971 UN Convention
on Pyschotropic Substances, their use is controlled by legislation.
For example, under Section 21 of the UK Drugs Act 2005, psilocybin
and its derivative psilocin are Class A drugs, and the knowing
possession of mushrooms that produce these, magic mush-
rooms, is illegal. There are, however, at least 30 magic mush-
rooms that grow in the UK, of which Psilocybe semilanceata is the
most commonly collected and used. The highest reported
concentrations of psilocybin are from P. azurescens (blue angels)
which is to be found in the Pacic north-west of the USA [89].
However, concentrations may vary widely within a single species
as a result of biological or ecological factors. Psilocybe cubensis, P.
mexicana, and P. semilanceata are the most-used species, but
worldwide, there are at least 216 mushrooms known to have
neurotropic effects; occurrences of these, listed by country, have
been compiled [91]. Over 150 of these fungi are species of Psilocybe
[90]; there are 230 or so species of this genus that are not easy to
identify without critical morphological and microscopical exam-
inations. The psilocybin-containing species generally turn bluish
on handling or bruising, especially on the stalks (stipes), but this is
not an unequivocal guide as various other fungi with different
compounds can give a similar reaction. Particular neurotropic
Psilocybe species can be encountered in forensic situations outside
their normal geographical areas as they are exported from Central
America into Europe, and some can be cultivated both indoors and
outside [92]. As some harmless species are supercially rather
similar to hallucinogenic ones, accurate identication is critical to
law enforcement. Interestingly, as a result of recent molecular
phylogenetic studies, many of the non-hallucinogenic species
traditionally classied in Psilocybe are now being placed under the
generic name Deconica [93]. Some species that contain the
prohibited compounds do not resemble Psilocybe species, for
example, the blue-green capped Stropharia aeruginosa. The
compounds psilocin and psilocybin can, however, be detected
by relatively inexpensive thin-layer chromatographic (TLC)
methods [94,95]. Procedures for the chemical determination of
psilocybin and psilocin by gas chromatography and mass
spectrometry (GCMS), gas liquid chromatography (GLC), high
performance liquid chromatography (HPLC), and spectroscopy
have also been documented [95,96]. Chemical methods are
especially valuable for testing magic mushroom preparations in
the form of powders, tablets, or capsules.
In view of the difculty in differentiating the hallucinogenic
Psilocybe species, several groups of researchers have endeavoured
to develop molecular tools for their discrimination. Lee et al. [97]
compared the ITS-1 region sequences of ve species and these
appeared useful at the generic level. However, a larger study
involving 35 North American Psilocybe species, and six other
genera, demonstrated that the rDNA ITS-1 region was too variable
to provide satisfactory resolutions and that nLSU rRNA gave better
correlations [98]. Using ITS and LSU sequences of four Psilocybe
species, Maruyama et al. [99] provided an improved identication
system. However, as there are over 200 species accepted in the
genus, not all of which are hallucinogenic (see above), further work
involving a more complete range of species is needed before the
molecular data will be robust enough to be used unequivocally in
court.
D.L. Hawksworth, P.E.J. Wiltshire / Forensic Science International 206 (2011) 111 7
While not normally fatal, irresponsible use of Psilocybe species
may lead to death either as a result of erratic behaviour while
under the inuence of the drugs they contain, or as a result of
imbibing extracts of the mushroom together with other drugs.
At present it is only psilocin and psilocybin-containing mush-
rooms that are prohibited from use by law in the UK. Other
pyschotropic mushrooms include the y agaric (Amanita muscaria)
which has the familiar scarlet white-ecked cap. This has been
used in religious ceremonies for centuries, has an inebriating
effect, and is not fatal itself unless consumed in large amounts.
With the prohibition of hallucinogenic Psilocybe species, dealers in
north London have been found selling the y agaric mushroom as
an alternative.
Some mould fungi also produce toxins directly (see below), or
have products that can react with other compounds to produce
toxic substances. Most notorious in this connection is Scopular-
iopsis brevicaulis. This fungus has been implicated in the
production of trimethyl arsine from arsenic-containing com-
pounds. This gas can be produced by the fungus in damp conditions
when growing on wallpaper containing Paris Green, and its
inhalation can lead to death [100]. In the early 1990s, it was
suggested that this same fungus reacted with arsenic-containing
re-retardants in cot mattresses to produce the toxic gas and that
this was the cause of Sudden Infant Death Syndrome (SIDS). While
that hypothesis did not stand up to critical examination [101],
other fungi occur in bedding materials, and some of these can
produce other toxins. Where there is incidence of suspicious death
in beds, ofcers attending scenes should check for any substantial
fungal growth.
7. Location of corpses
While we are not aware of any criminal case where fungi have
yet had a role in the locationof buried corpses, there might be some
potential. Some mushrooms are characteristic of disturbed ground,
but will not produce sporophores until 12 years after the
disturbance. Furthermore, some of the species with this ecology
are easy to recognise, notably the shaggy ink cap (Coprinus
comatus), and some morels (primarily Morchella species).
Since Sagara [102] reported the association between shallowly
buried carcasses of a dog and a cat and the mushroom Hebeloma
vinosophyllum in two different sites in Japan in 1976, there has
been considerable interest in the potential of corpse-nder fungi.
That species is known to appear 38 months after urea or
ammonium application to soils and then to continue to produce
mushroom sporophores for 12 years. The potential was empha-
sized by Carter and Tibbett [103,104], who distinguished between
an assemblage of ammonia fungi (AF) making sporophores after
the addition of urea or ammonia, and post-putrefaction fungi
(PPF) that form sporophores over animal cadavers without such
additions. Their paper was criticised by Bunyard [105] on the
grounds that of the 35 species they listed, only Hebeloma radicosum
and the North American Hebeloma syriense had been mentioned in
the literature as associated with corpses [106,107] rather than
ammoniacal substrates, and there were fewrst-hand claims. He
also stressed the problems of identifying these mushrooms, with
over 200 species of the genus known in North America alone, and
because some of the listed species had different ecologies. In
defending their position, Tibbett and Carter [108] accepted some of
Bunyards comments and emphasized that taphonomic mycota is
little more than a concept at this stage and requires further
research and development prior to practical application. That
remains the situation today, and we have not traced any case
where these fungi have assisted in nding, or been associated with,
a human grave. Hebeloma radicosum is widespread, but not very
common, in Western Europe; there are 187 records in the FRDBI
but none is associated with human remains, and no such records
are known to specialists in the genus (H. Becker, pers. comm.). A
detailed review of the association between fungi and graves, or
excrement of different organisms, has been prepared [57] and this
includes a list of 23 ammonia and post-putrefaction fungi, with
references to pertinent primary literature.
More practical in the location of corpses, are fungal indications
of where tree branches or logs have been disturbed or moved.
Mushrooms are generally geotropic,
11
with vertical stalks (stipes)
and horizontal caps (pilei), but if they are re-orientated, the stalks
may curve as they grow to regain a vertical position or their caps
skew to become horizontal once more [109]. In a case for South
Wales Police, this phenomenon was used to refute the statement of
the nder of a murder victims grave where he claimed he had not
touched or disturbed anything at the scene. Logs placed over the
surface of the grave by the offenders supported numerous
sporophores of a Marasmius species. The orientation of the stipes
and pilei showed that several of the logs had been turned over
some (indeterminate) time before the nd was reported [110] and
his testimony was refuted. If time were critical in such investiga-
tions, simple experiments could be conducted to test the length of
time required for reorientation of the fungus.
Lichens, particularly foliose ones, normally occur on the better-
illuminated sides of branches, so if they are found on an underside
of a broken branch then it is likely to have been disturbed. Further,
some lichens undergo colour changes before they decompose if, for
example they become dislodged or are covered (see above). This
phenomenon is most easily seen in shrubby and foliose species
containing b-orcinol depsidones such as norstictic or salazinic
acids (e.g. certain species of Parmelia and Usnea). These assume a
pinkred colour if moved to densely shaded or wet situations and
they gradually die [111,112]. Investigators should be sensitive to
such phenomena as they have the potential of indicating where
branches have been disturbed or moved. This may help identify
offender pathways at crime scenes, or recognition of clandestine
graves.
8. Biological warfare
While the possibilities for the use of bacteria and viruses in
biological warfare are well known [113], there is relatively little
appreciation of the potential of fungi in this respect. Many moulds
as well as mushrooms can produce toxins and, whilst the action of
most is long-term(for example they are carcinogens), some can be
cultured in vats in large amounts and produce quicker-acting
substances that have potential as biological weapons. They were
wrongly implicated in Yellow Rain in Vietnam, but more
importantly, within the last 30 years, a certain middle-eastern
country was found trying to purchase toxin-producing strains of
Fusarium from microbial culture collections in North America and
Europe. A review of the potential of toxin-producing fungi as
biological weapons has been published [114]; most dangerous is
the Fusarium T2 toxin.
Fungal parasites of plants can also be biological warfare agents
as they can be developed to destroy crops in the same way that
they are used for weed control [115]. The technology for spraying
spore suspensions of fungi from aircraft is now developed to a
sophisticated level a by-product of the use of Metarhizium
anisopliae var. acridum as a mycoinsecticide (Green Muscle) for
the control of locusts and grasshoppers [116]. If suspected
terrorists are found with what appear to be living cultures of
fungi, the material should be conscated and examined by a
specialist.
11
Affected by gravity.
D.L. Hawksworth, P.E.J. Wiltshire / Forensic Science International 206 (2011) 111 8
9. When to consider forensic mycology
In view of the foregoing, it is evident that the situations in
which investigating ofcers should consider utilizing mycology
can be summarised as follows:
(1) As an integral part of the ecological assessment at crime scenes,
especially in outdoor situations.
(2) When the time of death or deposition is uncertain, and fungal
colonies are evident on human remains, clothing, or associated
items (indoors or out of doors). This is particularly important if
entomology is not appropriate but, if fungi are present as well
as ies, fungi can be considered as an independent line of
evidence.
(3) If trace evidence is being sought and fungal spores are found in
palynological preparations.
(4) When mushrooms are found in the possessions of a suspect, in
gut contents, or in food and drink associated with deaths or
neurotropic behaviour.
(5) If fungi are being grown in mass-culture (e.g. in liquid growth
media in large containers).
10. Developing forensic mycology
While mycology has been demonstrated to provide useful
forensic evidence in a variety of ways and, in some cases, has
proved critical in securing convictions, it is currently rarely
employed. The principle reason for this is a lack of awareness
amongst crime scene investigators, and investigating ofcers, that
fungi have the potential to make signicant contributions. In
several of the cases in which we have been involved, it was the
astuteness of ofcers in recognising there was fungal growth
present on human remains or artefacts that led to our being called
in.
A secondary reason is a shortage of appropriately skilled
mycologists. Some categories of mycological investigation, partic-
ularly those involving the isolation and culture of fungi from
remains and materials, can be undertaken by biologists with
experience in microbiological methods, but those requiring
identication need specialists of wide experience. There is a
shortage of professional mycologists able reliably to identify fungi
across different systematic groups. For example, in the UK, there
has been a reduction in systematic mycology posts in the last 10
15 years as a result of the restructuring of research institutes and
retirements. The number of full-time fungal (including lichen)
systematists in the country (excluding curators of collections) is
currently a mere eight (of which two are lichenologists); and not
one is nowemployed anywhere in the university sector. This gure
of eight compares with 23 in 1997, and the House of Lords Science
and Technology Committee has considered the situation so grave
as to be generally recognised as a crisis [117]. The UKis not unique
in this respect, and action is needed to redress this problemso as to
ensure that adequate support is available not only for forensic
science, but also food safety, human health, plant and tree diseases,
pharmaceutical development, environmental assessments, and
nature conservation. No quick-x is possible as it takes at least a
PhD, plus 10 years experience, to produce a competent profes-
sional systematic mycologist.
The subject is also difcult because of a lack of comprehensive
identication manuals. This makes it hard for a non-specialist to
make critical identications. Although there are monographic
treatments of some groups, even to know which work to use
requires specialist knowledge. Because the number of species is
high and many are still unknown, this is a particular problem in
fungi, compared with, for example, plants. Furthermore, previously
unknown or unrecognised species are discovered every year. Semi-
popular guides rarely treat all species of a genus, and often do not
indicate that there are other similar species. There is not even a
pocket-sized identication manual on all hallucinogenic fungi,
something that would make the psilocybin and psilocin possession
section of the Drugs Act 2005 more practical for police ofcers to
enforce. Those undertaking aspects of forensic mycology that
involve numerous identications, as in the provision of trace
evidence, consequently require access to a substantial library and
to reference collections of authoritatively named material. The
most reliable way to identify fungi is by direct comparison with
correctly named reference material.
Identications using molecular sequence data can be made in
certain well-researched groups of fungi, especially those involved
in food spoilage, human diseases, or plant pathology [118]. The
existing DNA databases, however, have only a small proportion of
the known fungi represented in them, and many are represented
by only one or a few strains, so there is little representation of
within-species variability. Further, the gap between those known,
and those that are sequenced, may be growing; only 21% of the
species described in 200307 are represented in GenBank [119].
Also, and especially worrying, is that several independent studies
indicate that around 20% of the fungal sequences deposited in
GenBank are based on wrongly identied material [120]. This
means that while in some cases DNA data may be able to provide
conclusive identications, in many others either it will not, or it
will yield erroneous results. Specialist interpretation is, therefore,
essential in using DNA (e.g. BLAST) approaches for fungal
identication; this is not something that is yet at a stage where
it can be delegated to non-specialists.
Acknowledgements
We are grateful for the excellent assistance, support, and
expertise we have received from Dr. Judy Webb and Ms Julia
Newberry in our forensic case work. We would also like to thank
Dr. Jane Nicklin for her enthusiastic help, Professor Henry Becker
for information on Hebeloma species, and two anonymous
reviewers. Last, but not least, we gratefully acknowledge the
cooperationand support of ofcers frommany British Police forces.
In particular in relation to cases discussed here, we thank
Hertfordshire Constabulary, Lincolnshire Police, Metropolitan
Police Service, South Wales Police, Suffolk Constabulary, Sussex
Police, Tayside Police, Thames Valley Police, Wiltshire Constabu-
lary, and the British Transport Police. This work was completed
while D.L.H. was receiving support from the Ministerio de
Educacio n y Ciencia of Spain (Proyectos I+D CGL 2008-01600).
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