Hair identification key of wild and domestic ungulates from southern Europe

Abstract

We analysed macro- and microscopic features of dorsal guard hairs in 105 specimens of 10 wild and five domestic ungulates from southern Europe to work out a dichotomous key with a photographic reference system of diagnostic hair features. We integrated and extended the available data on hair morphology of wild ungulates and provide a first comparative analysis of hair structure of domestic forms. To develop the key, we used clearly recognisable qualitative characters of cuticle and medulla. The techniques used in this study can be easily, quickly and economically applied in routine investigations, keeping the time required to identify a sample at a minimum. The accuracy of the key was assessed through a blind test carried out by four trained observers. We describe the effects of age and season on the microscopic structure of hair, which have not yet been described in European literature. A review of all the available data on hair morphology of wild ungulates is presented and the relevant differences between domestic forms and their relative wild ancestors that have arisen during the domestication process are described. A hair identification key has a wide range of practical applications in biology, such as the study of carnivore feeding habits through scat analysis.

Full text

Hair identification key of wild and domestic ungulates from
southern Europe
Anna Maria De Marinis & Alessandro Asprea
De Marinis, A.M. & Asprea, A. 2006: Hair identification key of wild and domes-
tic ungulates from southern Europe. - Wildl. Biol. 12: 305-320.
We analysed macro- and microscopic features of dorsal guard hairs in 105 spe-
cimens of 10 wild and five domestic ungulates from southern Europe to work
out a dichotomous key with a photographic reference system of diagnostic hair
features. We integrated and extended the available data on hair morphology of
wild ungulates and provide a first comparative analysis of hair structure of domes-
tic forms. To develop the key, we used clearly recognisable qualitative characters
of cuticle and medulla. The techniques used in this study can be easily, quickly
and economically applied in routine investigations, keeping the time required
to identify a sample at a minimum. The accuracy of the key was assessed through
a blind test carried out by four trained observers. We describe the effects of age
and season on the microscopic structure of hair, which have not yet been de-
scribed in European literature. A review of all the available data on hair mor-
phology of wild ungulates is presented and the relevant differences between
domestic forms and their relative wild ancestors that have arisen during the
domestication process are described. A hair identification key has a wide range
of practical applications in biology, such as the study of carnivore feeding hab-
its through scat analysis.
Key words: bovids, deer, domestic, guard hairs, identification key, southern
Europe, suids
Anna Maria De Marinis, Istituto Nazionale per la Fauna Selvatica, Via Ca’
Fornacetta 9, I-40064 Ozzano dell’Emilia (BO), Italy - e-mail address: annamaria.
demarinis@infs.it
Alessandro Asprea, Servizio Scientifico, Abruzzo Lazio and Molise National
Park, Viale Santa Lucia, I-67032 Pescasseroli (AQ), Italy - e-mail address: ales
sandro.asprea@infinito.it.
Corresponding author: Anna Maria De Marinis
Received 23 November 2004, accepted 20 May 2005
Associate Editor: Klaus Hackländer
Identification of hair of mammalian species has practi-
cal applications in forensic medicine, taxonomy, palae-
ontology, zooarchaeology, anthropology and ecology.
In particular, the study of predator feeding habits from
the analysis of prey hairs found in scats has been wide-
ly used for describing the diet of mammalian carnivores,
because this technique is non-destructive and scats are
easy to collect throughout the year. In southern Europe,
several studies on the feeding habits of the wolf Canis
lupus have been carried out during the last two decades
(cf. Meriggi & Lovari 1996, Ciucci & Boitani 1998a).
Wild and domestic ungulates represent the main com-
ponent of wolf diet in relation to their local abundance
and/or local accessibility (Meriggi & Lovari 1996).
Predation on livestock is the crucial socio-economic fac-
tor promoting wolf persecution (cf. Ciucci & Boitani
1998b, Boitani 2003). Therefore the knowledge of wolf
diet plays a critical role in terms of conservation of one
of the last large carnivores in southern Europe. An iden-
tification key to the hairs of wild and domestic ungu-
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lates has a valid practical application in studies on wolf
feeding habits.
Several atlases (Faliu et al. 1980, Debrot et al. 1982)
and identification keys (Day 1966, Dziurdzik 1973, Kel-
ler 1978, 1980, 1981a, 1981b, 1992, Teerink 1991, Meyer
et al. 2002) have been published on European mammal
hairs. Only some of them deal with ungulates, but none
includes all the species occurring in southern Europe.
For example, the hairs of wild goat Capra aegagrus and
southern chamois Rupicapra pyrenaica have not been
described previously. The description of the hair struc-
ture of the different species is not always linked to
dichotomous keys (Debrot et al. 1982, Faliu et al. 1980).
Existing keys do not allow identification at the species
level (Dziurdzik 1973, Meyer et al. 2002) or concern
only a few species (Keller 1981b, Teerink 1991). More-
over the study of hair morphology of domestic ungu-
lates (Abad 1955, Dziurdzik 1973, Faliu et al. 1980, Kel-
ler 1992) is not exhaustive. Only Keller (1992) com-
pared wild equids with domestic forms. The effects of
age and season on hair structure of ungulates have not
been investigated in these studies. Last, but not least,
each of these manuals is based on different techniques,
characteristics and nomenclature. Since in many cases
it is necessary to consult and compare several works at
once, some confusion can arise during identification.
Moreover, some of these studies require special tech-
niques and equipment such as scanning electron micro-
scopes and microtomes which, in turn, require skilled
labour, thus slowing down the analysis.
The aims of our paper are 1) to integrate and extend
the available data on hair morphology of wild ungulates
occurring in southern Europe, 2) to present the first com-
parative analysis of hair structure of domestic forms, and
3) to provide an unambiguous discrimination tool be-
tween hairs of ungulates, based on a dichotomous key
and a photographic reference system.
Material and methods
We studied the hair structure of 10 wild ungulate spe-
cies: wild boar Sus scrofa, fallow deer Dama dama, red
deer Cervus elaphus, roe deer Capreolus capreolus,
southern chamois (Apennine population) Rupicapra pyre-
naica, alpine chamois R. rupicapra, alpine ibex Capra
ibex, Spanish ibex Capra pyrenaica, wild goat Capra
aegagrus and mouflon Ovis orientalis. Animals sam-
pled came from the Italian peninsula, Sardinia and
Corsica, except for Capra pyrenaica (Spain) and Capra
aegagrus (Montecristo Island and Crete). We compared
the hair structure of these wild ungulates with that of 12
breeds and cross-breeds of five domestic ungulates (cow,
sheep, goat, horse and donkey). Our identification key
does not include ungulate species recently introduced in
southern Europe which have only formed very small
populations at geographically restricted ranges, such as
e.g. axis deer Axis axis occurring in the Brijuni Islands,
Croatia (Mitchell-Jones et al. 1999). The names of wild
species follow Wilson & Reeder (1993), and the taxon-
omy has not been updated to be in line with the new
accounts prepared for Wilson & Reeder (2005) to make
easier the analysis of our review and the comparison
with previous works. Terrestrial mammals are covered
with two distinct types of hairs: long, thick, pigmented
guard hairs, determining the general colour of the coat,
and short, thin, less pigmented and more numerous fine
hairs, supplying insulation. Only guard hairs are impor-
tant in species identification as they exhibit diagnostical-
ly reliable features. Since the hair structure of fresh and
tanned specimens is identical (Mayer 1952, Perrin &
Campbell 1980, Hess et al. 1985), we collected hair tufts
from live or freshly hunted animals, and dry skins housed
in Italian mammal collections (Natural History Museum,
Zoological Section, Florence; Italian Wildlife Institute,
Bologna; Regional Museum of Natural Science, Chateau
de Saint-Pierre). Hair samples were taken from 105 spe-
cimens (75 wild and 30 domestic). The hairs were col-
lected from not less than five spots in the dorsolateral
body region of each specimen. Hairs of other body
regions generally show similar characteristics, but they
are often less marked and thus hardly identifiable (Amera-
singhe 1983, Teerink 1991). We did not include in this
key specialised kinds of guard hairs, i.e. bristles. Previous
studies reported that the first moult in young cervids and
bovids of some European and North American species
coincides with the development of adult hair character-
istics (cf. Scott & Shackleton 1980, Kennedy & Carbyn
1981, Jedrzejewski et al. 1992, Gade-Jørgensen & Stage-
gaard 2000). As far as we know, no data are available
on domestic ungulates. To study the effect of age on hair
structure of wild and domestic ungulates, we defined
two age classes: young (prior to the first moult) and
adult. We sampled 28 young of known age (< 4 months
for wild species, < 2 months for domestic forms) and 69
adults. To study the effect of season on hair structure of
wild ungulates, we collected hair tufts from winter (N =
25) and summer (N = 15) coats.
Pigmentation and hair dimension are variable charac-
ters and their usefulness in an identification key is cor-
respondingly limited. They change with age, season and
body region. Moreover, pigmentation can be deteriorat-
ed, e.g. under the action of the digestive enzymes, while
hair dimension can be modified, e.g. because of frag-
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mentation during digestion (Kennedy
& Carbyn 1981, Amerasinghe 1983).
Only hair profile and microstructure
have diagnostic value and are species
representative. The guard hairs of un-
gulates do not show any expanded and
flattened regions (Kennedy & Carbyn
1981). We distinguished two general
regions along the hair, each one cor-
responding to half a hair, i.e. the low-
er and upper shaft (Moore et al. 1974).
Hair microstructure is described refer-
ring to these regions, firstly because
microstructure changes along the hair
(Wallis 1993, Oli 1993) and lastly be-
cause hairs can be fragmented and only
lower or upper shafts can be exam-
ined.
Taking into account the nomencla-
ture currently used to describe hair
profile and microstructure (cf. Kenne-
dy & Carbyn 1981, Teerink 1991), we
selected only a few descriptive cate-
gories, easily identifiable without mis-
understandings. We distinguished two
types of hair profile: undulated and
straight. The hair tip is described as
split or not split. Hair microstructure
is composed of three layers of kera-
tin: medulla, cortex and cuticle, from
the innermost to the outermost. The
medulla is composed of loosely packed
cells with air spaces in the cells them-
selves or between them. It is described
(Fig. 1) by composition (unicellular
irregular and multicellular), structure
(amorphous, uniseriate, multiseriate,
vacuolated, filled lattice and partially
filled lattice), pattern (continuous and
fragmental) and margin form (irregu-
lar, straight and scalloped). The cor-
tex, composed of cells coalesced into
a hyaline mass, does not exhibit char-
acters which could be used as criteria
for identification because of its near-
ly homogeneous structure. However,
there is a considerable interspecific
variation in the cortex width and thus
we included this parameter in the key.
The cortex width was estimated by
eye, taking the width of the medulla
as a reference unit. The cuticle con-
Compo-
sition
Unicellular irregular Multicellular
Structure
Amorphous Uniseriate Multiseriate Filled lattice Partially filled lattice Vacuolated
Pattern
Continuous Fragmental
Form of the
margins
Irregular Straight Scalloped
Figure 2. Cuticle classification system used in our hair key to identify wild and domestic
ungulates from southern Europe.
Figure 1. Medulla classification system used in our hair key to identify wild and domestic
ungulates from southern Europe.
Position
Transversal Intermediate
Structure of
scale margin
Smooth Rippled Heavily
Distance
between
scale margin
Distant Near
Scale pattern
Regular mosaic Regular wave Irregular wave 7shaped
Slightly
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sists of overlapping scales. We considered (Fig. 2) the
position of the scales in relation to the longitudinal axis
of the hair (transversal and intermediate), the structure
of scale margins (smooth and rippled), the distance
between scale margins (distant and near) and the scale
pattern (regular mosaic, regular and irregular wave, 'Ω'-
shaped).
Microscopic hair preparations were made following
De Marinis & Agnelli (1993). Hair cross-sectioning is
the most complicated step of the whole microscopic
analysis. The shapes of cross-sections change along the
hair and only the sequence of the shapes along the hair,
rather than the shape at any particular point, is impor-
tant for the identification (Teerink 1991). Furthermore,
cross-sections of hairs of closely related species, such
as those included in this key, appear very similar to each
other, without diagnostic value (Kennedy & Carbyn
1981). Practicability is an important factor to take into
account when selecting diagnostic criteria, thus hair
cross-sections have not been included in this key. Micro-
photographs were taken using a light microscope fitted
with a digital camera. Only microphotographs of repre-
sentative and predominant patterns were included in the
key in order to provide a valid help during the succes-
sive identification steps (see microphotographs in Appen-
dix I).
The accuracy of the key developed in this study was
assessed through a blind test on a sample of 260 hairs
randomly taken from our reference collection. Four
trained observers identified 65 samples each, to the lev-
el of species, age class and season.
Results and discussion
Macroscopic hair description
Among wild ungulates, only wild boar hairs can be sure-
ly identified without the aid of microscopic analysis.
The hair of the wild boar can be easily identified by eye
on the basis of the general appearance of the hair, which
is split at least once. Piglets do not show split tips. How-
ever, they have bristly hairs which can be distinguished
from those of other ungulates even if the observer is not
highly skilled in hair identification (Table 1). Frayed
guard hairs also characterise domestic and feral swine
and hybrids (Marchinton et al. 1974, Hess et al. 1985,
Mayer & Brisbin 1991). Boar subspecies and other
Suidae species have bristles with typically frayed ends
(Koppiker & Sabnis 1977, Amerasinghe 1983, Hess et
al. 1985). Moreover, the Tayassuidae species, ecologi-
cal equivalents of suids in the New World, have guard
hairs split at the tip (Hess et al. 1985). Degree of fray-
ing might be correlated with chronology of hair replace-
ment (Hess et al. 1985). The adaptive significance of
this character of hair morphology is not known. Adults
of cervids and wild bovids can be macroscopically dis-
tinguished from domestic forms because of their shiny
appearance and undulated profile (see Table 1). Hairs
of young other than sheep can be distinguished from
those of adults because they are thin and straight, in wild
and domestic forms as well (see Table 1). However, young
of wild species have a shiny appearance as do adults.
Adults and young of sheep show undulated and dull hairs
(see Table 1). The macroscopic analysis of the guard
hairs represents only the first step in the identification
process of an unknown hair sample. Moreover, the mac-
roscopic analysis can hardly be carried out when exam-
ining fragmented hairs, such as those found in carnivore
scats.
Microscopic hair description
Medulla features
We identified two different medullary structures in wild
ungulates: amorphous in wild boar and lattice in the oth-
ers (Table 2). We did not find the medullary lattice struc-
ture in wild boar hairs, as previously reported by Faliu
et al. (1980) and Keller (1981b). This structure has been
observed by Amerasinghe (1983) only in the bristles of
the mid-dorsal line of the wild pig in Sri Lanka, howev-
er, it was much reduced and fragmental. The medulla
structure per species is constant throughout the hair
length, and it does not change with age or season. A
strong structural homogeneity in medullary pattern can
be observed in the whole Cervidae family throughout the
world. A review of 21 studies carried out on the micro-
scopic features of deer hairs revealed that all the species
have a medullary structure which is filled lattice (Appen-
dix II). As far as it is currently known, a similar struc-
Table 1. Macroscopic characters distinguishing hairs of wild and domestic ungulates.
Ungulate Family Age class Profile General appearance Tip
Wild Suidae Adult and piglet Straight Bristly Split, except in piglet
Cervidae/Bovidae Adult Undulated Thick and shiny Not split
Cervidae/Bovidae Young Straight Thin and shiny Not split
Domestic Bovidae/Equidae Adult and young Straight (undulated in sheep) Thick (thin in young) and dull Not split
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tural homogeneity is found in bovids but only at the sub-
family level. For example, re-examining 17 studies on
the hairs of Caprinae and Bovinae, the former shows a
medullary lattice structure while the latter presents a
multiseriate structure (Appendix III). Different patterns
of medullary cells might express the evolutionary his-
tory of the species (Sheng et al. 1993, Chernova 2001),
although ecological factors almost certainly play an
important role in determining hair morphology (cf. Perrin
& Campbell 1980).
The domestic ungulates showed multiseriate and uni-
seriate medullary structures (Table 3). Sheep and goats
have a medullary structure that differs from that of their
relative wild ancestors, i.e. Ovis orientalis and Capra
aegagrus, respectively. During the domestication pro-
cess important structural modifications in the architec-
tonics of medulla were induced by artificial selection as
affected by economic, cultural and aesthetic reasons
(Clutton-Brock 1999, Meyer et al. 2002). These varia-
tions are especially prominent in mammals that have
undergone a long domestication process, such as sheep
and goats. Differences in hair structure are less obvious
in species that have been bred less intensively (Meyer et
al. 2002). Our hair samples of Ovis orientalis and Capra
aegagrus came from animals living respectively on
Corsica and Sardinia, and on Montecristo and Crete.
Generally speaking, Mediterranean islands often repre-
sent natural enclosures where goats and sheep have been
kept and bred since prehistory in a free ranging state
(Masseti 1998). Therefore our samples are not represen-
tative of the true wild ancestor of sheep and goats; they
come from relic populations of animals that were intro-
duced on the Mediterranean islands and now live as wild
populations (Clutton-Brock 1999, Masseti 2002). Never-
Table 2. Medulla features of wild ungulates according to age class.
Age class Taxon Medulla Cortex widthComposition Structure Pattern Form of the margins
Adult Sus scrofa Cells not clearly
visible Amorphous Continuous, fragmental
at the base Irregular, especially
in the upper shaft Wider than medulla in
the lower shaft, more
and more narrow in
the upper shaft
Dama dama
Cervus elaphus
Capreolus capreolus Multicellular Filled lattice with
polygonal cells
Continuous, suddenly
interrupted at the base Medulla fills
the entire
width of the hair
Very narrow or not
visible
Rupicapra pyrenaica
R. rupicapra
Ovis orientalis
Continuous, fragmental
at the base
Capra ibex
C. pyrenaica
C. aegagrus Multicellular Partially filled lattice
with elongated cells Continuous, fragmental
at the base Scalloped Clearly visible
Young Sus scrofa As in the adult As in the adult
Dama dama
Cervus elaphus
Capreolus capreolus Multicellular Partially filled lattice
with polygonal cells Continuous, fragmental
at the base Scalloped Clearly visible
Rupicapra pyrenaica
R. rupicapra
Ovis orientalis
Capra ibex
C. pyrenaica
C. aegagrus As in the adult As in the adult
Table 3. Medulla features of young and adult domestic ungulates.
Domestic form
Medulla
Cortex widthComposition Structure Pattern Form of the margins
Cow Multicellular Multiseriate, appearing
vacuolated Continuous Straight or irregular Wide as medulla or ½,⅓ of medulla
Goat Multicellular and
unicellular irregular Multiseriate and
uniseriate Continuous or
fragmental Scalloped and
irregular Very narrow but well recognisable if medulla is
multicellular, wider than medulla if medulla is
unicellular
Sheep Multicellular Multiseriate Continuous Scalloped Very narrow, but well recognisable
Horse Multicellular and
unicellular irregular Multiseriate and
uniseriate Continuous or
fragmental Scalloped and
irregular ½ of medulla if medulla is multicellular, wider
than medulla if medulla is unicellular
Donkey Multicellular Multiseriate Continuous Straight and irregular Very narrow, but well recognisable
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theless, the medullary structures are strongly different
between these 'wild' forms and their domestic counter-
parts, because domesticated animals that return to the
wild, will usually revert by natural selection to a physical
form that is closer to the wild species (Clutton-Brock
1999). In fact Capra aegagrus and Ovis orientalis kept
the identical medullary structure of the relative species
of the same genus (see Table 2) which have never been
domesticated (Clutton-Brock 1999).
Cuticle features
We did not observe a clear differentiation in the cuticle
features between domestic and wild ungulates, as found
for the medulla (Tables 4 and 5). The cuticle was a use-
ful tool to discriminate species among wild ungulates
and to distinguish young from adult and winter from
summer coat in deer (see Table 4). Meyer et al. (2001)
already differentiated three subfamilies within Cervidae
in a comparative structural analysis of hair cuticular pat-
tern. In their study, the subgroup differentiation was
obtained by calculating quantitative parameters such as
the ratio of scale width to scale height. These groupings
corroborated the zoosystematical relationships that have
been achieved using modern molecular genetic tech-
niques.
We could not use cuticle to distinguish domestic ungu-
lates from each other, because of high overlap among
cuticular patterns (see Table 5). Domestication induced
changes in several cuticular features of the hair shaft,
and the resulting structural homogeneity in the cuticle
compromises species identification ( Meyer et al. 1997,
2000). Sheep have a cuticular structure different from
their relative wild ancestors, i.e. Ovis orientalis, while
goats do not show clear differences in the cuticle from
their relative wild ancestors, i.e. Capra aegagrus (see
Tables 4 and 5). According to Meyer et al. (2001) there
is a relationship between the cuticle scale parameters and
the coat structure and function. Therefore, the strong arti-
Table 4. Cuticle features of wild ungulates according to age class and season.
Age class Taxon Season
Lower shaft Upper shaft
Scale
position Scale
margin Scale mar-
gin distance Scale
pattern Scale
position Scale
margin Scale mar-
gin distance Scale
pattern
Adult Sus scrofa Transversal Smooth Near Regular wave Transversal Heavily rippled Near Regular wave
Cervus
elaphus Winter Transversal
Smooth
Distant
Regular wave
Transversal
Generally smooth Distant Regular wave
Summer Generally
rippled Distant or
near
Capreolus
capreolus Winter Intermediate Smooth Distant Regular
mosaic Transversal Slightly
rippled Distant Regular wave
Summer Transversal Regular wave
Dama dama Winter Transversal
Smooth
Distant
Regular
wave Transversal
Generally slightly
rippled Distant Regular wave
Summer Slightly or
heavily rippled Distant or
near
Rupicapra pyrenaica
R. rupicapra
Capra ibex Transversal Smooth Distant Irregular
wave Transversal Generally
rippled Distant Regular wave
C. aegagrus
C. pyrenaica Transversal Smooth Distant Irregular
wave Transversal Generally
rippled Distant Regular wave
and 'Ω'-shaped
Ovis orientalis Intermediate Smooth Distant Regular
mosaic Transversal Generally
rippled Distant Regular wave
Young
Sus scrofa Transversal Smooth Distant Regular wave Transversal Rippled Distant Regular wave
Cervidae
Bovidae Transversal Smooth Distant Irregular
wave Transversal Smooth Distant Irregular wave
Table 5. Cuticle features of domestic ungulates.
Domestic form
Lower shaft Upper shaft
Scale
position Scale
margin Scale margin
distance Scale
pattern Scale
position Scale
margin Scale margin
distance Scale
pattern
Cow Transversal Smooth Distant Regular and irregular wave Transversal Smooth and rippled Distant Regular wave
Goat Transversal Smooth Distant Irregular wave Transversal Smooth and rippled Distant and near Regular wave
and 'Ω'-shaped
Sheep Transversal Smooth Distant Irregular wave Transversal Generally rippled Distant Irregular wave
and 'Ω'-shaped
Horse Transversal Smooth Distant Irregular wave Transversal Generally smooth Distant Regular wave
Donkey Transversal Smooth Distant Regular wave Transversal Rippled Near Regular wave
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ficial selection for wool production may have transformed
the hair structure of the sheep but not that of the goat.
Hair identification key
Our key (Table 6) is based on three main steps: 1) macro-
scopic analysis permitting the separation of adult wild
boar from all other ungulates; 2) analysis of medulla and
cortex features allowing the separation between wild
species and domestic forms, and the identification of
domestic ungulates; 3) analysis of cuticular features in
wild ungulates leading to the determination of age class
and species, and the distinction between winter and sum-
mer coats.
Keller (1981b) separated deer from wild bovids on the
basis of cross-sections. We distinguished these taxa by
the shape of the basal part of the hair (see Appendix I,
App. Figs. 10 and 11), without hair cross-sectioning,
which is a complicated and time consuming step of the
laboratory procedures.
In a few cases it was not possible to identify to spe-
cies level. The cuticular features of the species belonging
to the genus Rupicapra are practically identical. Difficulty
was encountered in differentiating the species of the
genus Capra; only Capra ibex can be separated on the
basis of the scale pattern of the upper shaft. However,
the range of these species does not overlap at all in south-
ern Europe (Mitchell-Jones et al. 1999).
The variability observed in the hair cuticular features
of the winter and summer coats of deer makes the spe-
cies identification more complicated. The species can
be separated from each other only when comparing win-
ter hairs. In summer, the differences between species are
less pronounced, and only fallow deer hairs are recog-
nisable. 'V' shaped incisions in the cuticular pattern of
the upper shaft have been reported as a diagnostic fea-
ture to distinguish roe deer from other deer (Lomuller
1924, Keller 1981b, Teerink 1991). We frequently observed
these 'V' shaped incisions in almost all wild and domes-
tic ungulates (see Appendix I, App. Fig. 1A), so they
should not be considered a reliable character for hair
identification.
We observed cylindrical root shape (see Appendix I,
App. Fig. 23) in horses and donkeys (90 and 82.6%
respectively, N = 53), and conical root shape (see Ap-
Table 6. Key to identifying wild and domestic ungulates from southern Europe based on hair samples. The figure numbers given below refer to Fig. 1 - Fig.
24 in Appendix I.
1. Hairs bristly, split at tip usually more than once .................................................................... Sus scrofa (adult)
Hairs other than above ......................................................................................................2
2. Medullary structure amorphous (Fig. 2) ...........................................................................Sus scrofa (piglet)
Medullary structure other than above ...........................................................................................3
3. Medullary structure lattice (Figs. 3 and 4) .......................................................................................4
Medullary structure other than above ...........................................................................................5
4. Medullary cells polygonal, practically filling the entire width of the hair so that the cortex is very narrow or
not visible (filled structure) (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
Medullary cells polygonal or elongated, not filling the entire width of the hair so that the cortex is clearly visible
(partially filled structure) (Fig. 4) ..............................................................................................7
5. Medullary structure vacuolated (Fig. 5) ......................................................................................cow
Medullary structure other than above (Figs. 6, 7, 8 and 9)...........................................................................8
6. Basal part of the hair wine-glass shaped (Fig. 10) .................................................................................9
Basal part of the hair slightly tapered (Fig. 11) ..................................................................................10
7. Cuticular scale pattern irregular wave and scale margins smooth throughout the hair length (Fig. 12) . . . . . . . . . . . . . . . . . . . .Young of deer and bovids
Cuticular scale pattern irregular wave and scale margins smooth only in the lower shaft..................................................11
8. Medullary composition only multicellular (Figs. 6, 7 and 8) ........................................................................12
Medullary composition multicellular and unicellular irregular (Fig. 9) ................................................................13
9. Cuticular scale pattern regular mosaic in the basal part of the lower shaft of the hair (Fig. 13) .......................Capreolus capreolus (winter)
Cuticular scale pattern regular wave throughout the lower shaft of the hair (Fig. 14) .....................................................14
10. Cuticular scale pattern regular mosaic in the basal part of the lower shaft of the hair (Fig. 15) ...................................Ovis orientalis
Cuticular scale pattern irregular wave throughout the lowershaft of the hair (Fig. 16) .......................................... Rupicapra sp.
11. Cuticular scale pattern generally regular wave in the upper shaft of the hair (Fig. 17) .............................................Capra ibex
Cuticular scale pattern generally 'Ω'-shaped in some tracts of the upper shaft of the hair (Fig. 18) .............Capra pyrenaica and Capra aegagrus
12. Medulla margins scalloped (Fig. 6) ........................................................................................sheep
Medulla margins straight and irregular (Fig. 7) ..............................................................................donkey
13. Very narrow cortex, where the medulla has a multicellular composition (Fig. 6); cuticular scale pattern
'Ω'-shaped in some tracts of the upper shaft of the hair (Fig. 19)...................................................................goat
Cortex ½ of medulla, where the medulla has a multicellular composition (Fig. 8); cuticular scale pattern
other than above .......................................................................................................horse
14. Cuticular scale margins generally smooth in the upper shaft of the hair (Fig. 20)......................................Cervus elaphus (winter)
Cuticular scale margins generally rippled in the upper shaft of the hair (Fig. 21) ........................................................15
15. Cuticular scales margins also heavily rippled and near in the upper shaft of the hair (Fig. 22)............................. Dama dama (summer)
Cuticular scale margins never heavily rippled in the upper shaft of the hair
........................................................Cervus elaphus (summer), Capreolus capreolus (summer), Dama dama (winter)
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312 © WILDLIFE BIOLOGY · 12:3 (2006)
pendix I, App. Fig. 24) in sheep and goats (90.5 and
100%, respectively, N = 53). It is not common to find
hairs with roots in the scats; when it happens, the root
shape can help in hair identification of goats and horses
(see point 13 of the key in Table 6).
Young of deer and wild bovids can be distinguished
from adults because they have an irregular wave cutic-
ular scale pattern throughout the hair length (see Table
4). All the hair samples of young animals that we exam-
ined showed this feature from birth until 3-4 months of
age. Young change their birth coat into a coat with hairs
similar to those of adults during their first moult (cf.
Ryder 1960, Johnson & Hornby 1980, Jedrzejewski et
al. 1992). Young of the domestic forms showed the same
microscopic hair features as adults.
We did not observe relevant differences within the
breeds and cross-breeds examined.
Macroscopic and microscopic features used in this key
allowed a correct identification with an accuracy that aver-
aged 97.7 ± 0.9% at the species level (in wild ungulates:
98.1%, N = 157; in domestic ungulates: 99%, N = 98),
100% (N = 154) at the age class level and 96.6% (N =
29) in the recognition of winter and summer coat of deer.
The misidentifications concerned mostly (50%) the
genus Capra. Generally, the mistakes occurred in the
identification of the 'Ω'-shaped cuticular pattern, the dis-
tinction between slightly and heavily rippled cuticular
patterns and the recognition of the partially filled struc-
ture of medulla. The experience of the observer affect-
ed the correct identification of the cuticular patterns. The
possible degradation of strongly pigmented hairs, occur-
ring during the depigmentation process, may lead to the
misidentification of the medullary structure.
Conclusion
Our dichotomous key allows hair identification based
on characters which are clearly recognisable, consider-
ing the effects of age and season on hair structure. The
techniques used in our study can be easily, quickly and
economically applied in routine investigations keeping
the time required to identify a sample at a minimum, but
yielding accurate identifications.
However, some suggestions have to be kept in mind
before using our key:
• Prepare a reference collection including samples of all
the species that are going to be investigated. This is par-
ticularly recommended when examining domestic
ungulates.
• Work preliminarily with known samples taken from
the reference collection until good confidence with the
identification criteria is gained. Use blind tests to deter-
mine proficiency of identification.
• Analyse several tufts of hairs of the same individual
because of the considerable variety of hair types
encountered in different body regions and even within
the same body region (see Appendix I, Fig. 1). None-
theless no general statement can be made on the num-
ber of hairs that constitute an adequate sample for iden-
tification (Mayer 1952, Day 1966, Meyer et al. 2002).
• Compare always a set of features and do not rely on a
single character, considering the appreciable degree of
interspecies overlap in certain characteristics.
Acknowledgements - we wish to thank the following curators
of the museum collections, colleagues and friends who kind-
ly supplied hair samples for this study: P. Agnelli (Natural
History Museum, Zoological Section, Florence University);
S. Busatta, N. Canetti, L. Carnevali, B. Franzetti, A. Monaco
and P. Montanaro (Italian Wildlife Institute, Bologna); I. Gri-
mod (Regional Museum of Natural Science, Chateau de Saint-
Pierre); A. De Santis, P. Di Pirro, P. Genov, G. Lippa, S.
Monti, D. Pasut, M. Piccin, C. Pizzutto and L. Zorn. Special
thanks are extended to all the staff of Servizio Scientifico of
the Abruzzo Lazio and Molise National Park with particu-
lar reference to L. Gentile, R. Latini and C. Sulli for techni-
cal and logistical assistance, and V. Mastrella for his valu-
able help with photography. We are sincerely grateful also to
F. Amato, G. Del Greco and M. Manca for their contribution
in different phases of the work. A. De Faveri (Italian Wildlife
Institute, Bologna) made the tables on hair morphology. We also
wish to thank P. Genovesi (Italian Wildlife Institute, Bologna),
M. Masseti (Department of Animal Biology and Genetics,
Florence University), P. Agnelli (Natural History Museum,
Zoological Section, Florence University) and two anonymous
reviewers who all provided critical and helpful suggestions that
helped improve the manuscript.
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Appendix I
Microphotographs of representative and predominant patterns used in our hair key to identify wild and domestic
ungulates from southern Europe.
Appendix Figure 1. Cervus elaphus (adult), variation of cuticular scale
pattern in the upper shaft of the hair in the same individual (400x); ‘V’
shaped incisions are visible in A.
Appendix Figure 2. Sus scrofa (piglet), medulla; upper shaft of the
hair (400x).
Appendix Figure 3. Cervus elaphus (adult), medulla; upper shaft of
the hair (100x).
Appendix Figure 4. Capra ibex (young), medulla; upper shaft of the
hair (400x).
Appendix Figure 5. Cow (adult), medulla; lower shaft of the hair
(400x).
Appendix Figure 6. Goat (adult), medulla; upper shaft of the hair
(400x).
A B
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Appendix Figure 7. Donkey (adult), medulla; upper shaft of the hair
(400x).
Appendix Figure 8. Horse (adult), medulla; upper shaft of the hair
(400x).
Appendix Figure 9. Horse (adult), medulla; upper shaft of the hair
(400x).
Appendix Figure 10. Capreolus capreolus (adult); basal part of the
hair (100x).
Appendix Figure 11. Rupicapra pyrenaica (adult); basal part of the
hair (100x).
Appendix Figure 12. Capreolus capreolus (young); cuticle, upper shaft
of the hair (400x).
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Appendix Figure 13. Capreolus capreolus (adult, winter); cuticle, lower
shaft of the hair (400x).
Appendix Figure 14. Dama dama (adult), cuticle; lower shaft of the
hair (400x).
Appendix Figure 15. Ovis orientalis (adult), cuticle; lower shaft of
the hair (400x).
Appendix Figure 16. Rupicapra pyrenaica (adult); cuticle, lower shaft
of the hair (400x).
Appendix Figure 17. Capra ibex (adult), cuticle; upper shaft of the
hair (400x).
Appendix Figure 18. Capra aegagrus (adult); cuticle, upper shaft of
the hair (400x).
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318 © WILDLIFE BIOLOGY · 12:3 (2006)
Appendix Figure 24. Goat (adult); root shape (100x).
Appendix Figure 19. Goat (adult), cuticle; upper shaft of the hair
(400x).
Appendix Figure 20. Cervus elaphus (adult, winter); cuticle, upper
shaft of the hair (400x).
Appendix Figure 21. Dama dama (adult), cuticle; upper shaft of the
hair (400x).
Appendix Figure 22. Dama dama (adult, summer); cuticle, upper shaft
of the hair (400x).
Appendix Figure 23. Horse (adult); root shape (100x).
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Appendix II
Review of 21 studies on medullary structure of Cervidae. All the species listed have a filled lattice structure.
Classification according to Wilson & Reeder (1993).
Subfamily Species Reference
Cervinae Axis axis Lomuller (1924), Amerasinghe (1983), Chehébar & Martin (1989),
Mukherjee et al. (1994)
A. porcinus Amerasinghe (1983), Sheng et al. (1993)
Cervus albirostris Sheng et al. 1993
C. elaphus Present study, Lomuller (1924), Jullien (1930), Lochte (1938), Mayer
(1952), Dziurdzik (1973), Moore et al. (1974), Kennedy & Carbyn
(1981), Keller (1981b), Debrot et al. (1982), Chehébar & Martin
(1989), Teerink (1991), Sheng et al. (1993), Meyer et al. (2002)
C. eldii Sheng et al. (1993)
C. nippon Sheng et al. (1993), Meyer et al. (2002)
C. unicolor Koppiker & Sabnis (1976), Amerasinghe 1983, Sheng et al. (1993),
Mukherjee et al. (1994)
Dama dama Present study, Jullien (1930), Lochte (1938), Dziurdzik (1973), Debrot
et al. (1982), Chehébar & Martin (1989), Teerink (1991), Meyer et al.
(2002)
Elaphurus davidianus Sheng et al. (1993)
Hydropotinae Hydropotes inermis Sheng et al. (1993)
Muntiacinae Elaphodus cephalophus Sheng et al. (1993)
Muntiacus crinifrons Sheng et al. (1993)
M. feae Sheng et al. (1993)
M. muntjak Amerasinghe (1983), Sheng et al. (1993)
M. reevesi Sheng et al. 1993)
Capreolinae Alces alces Jullien (1930), Lochte (1938), Dziurdzik 1973, Moore et al. (1974),
Kennedy & Carbyn (1981), Debrot et al. (1982), Sheng et al. (1993),
Wallis (1993), Meyer et al. (2002)
Capreolus capreolus Present study, Lomuller (1924), Jullien (1930), Lochte (1938),
Dziurdzik (1973), Faliu et al. (1980), Keller (1981b), Debrot et al.
(1982), Teerink (1991), Sheng et al. (1993), Meyer et al. (2002)
Hippocamelus antisensis Vazquez et al. (2000)
H. bisulcus Chehébar & Martin (1989)
Mazama americana Vazquez et al. (2000)
M. guazoubira Vazquez et al. (2000)
Odocoileus hemionus Mayer (1952), Moore et al. (1974), Kennedy & Carbyn (1981)
O. virginianus Mayer (1952), Moore et al. (1974), Kennedy & Carbyn (1981),
Tumlison (1983), Wallis (1993)
Pudu puda Chehebar & Martin (1989), Feder & Arias (1992)
Rangifer tarandus Lomuller (1924), Jullien (1930), Kennedy & Carbyn (1981), Debrot et
al. (1982), Sheng et al. (1993)
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Appendix III
Review of 17 studies on medullary structure of Caprinae and Bovinae. All the species listed have a structure which
is either lattice (Caprinae) or multiseriate (Bovinae). Classification according to Wilson & Reeder (1993).
Subfamily Species Reference
Caprinae Capra aegagrus Present study
C. caucasica Jullien (1930)
C. ibex Present study, Lomuller (1924), Jullien (1930), Mandelli (1960),
Couturier (1962), Keller (1981b), Debrot et al. (1982), Meyer et al.
(2002)
C. pyrenaica Present study, Jullien (1930)
Hemitragus jemlahicus Oli (1993)
Oreamnos americanus Jullien (1930), Moore et al. (1974), Kennedy & Carbyn (1981)
Ovis canadensis Mayer (1952), Moore et al. (1974), Kennedy & Carbyn (1981)
O. dalli Kennedy & Carbyn (1981)
O. orientalis Present study, Lomuller (1924), Jullien (1930), Lochte (1938),
Dziurdzik (1973), Debrot et al. (1982), Teerink (1991), Meyer et al.
(2002)
Pseudois nayaur Oli (1993)
Rupicapra rupicapra Present study, Jullien (1930), Lochte (1938), Couturier (1938),
Mandelli (1960), Dziurdzik (1973), Faliu et al. (1980), Keller
(1981b), Meyer et al. (2002)
R. pyrenaica Present study
Bovinae Bison bison Mayer (1952), Moore et al. (1974), Kennedy & Carbyn (1981)
B. bonasus Dziurdzik (1978), Meyer et al. (2002)
Boselaphus tragocamelus Mukherjee et al. (1994)
Tetracerus quadricornis Mukherjee et al. (1994)
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