I

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Hans Brunner and Brian Coman

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Contents

Foreword

(likata Press Proprietary Limited

Melbourne

vii

Preface ix

First puhlished 1974

Acknowledgements

H. BRUNNER & B. COMAN

ix

SECTION A
National Library of Australia Card Number
and ISBN 0 909605 01 7

1. Introduction

General
Types of hair 2
The hair profile 3
Aspects of hair growth

Library of Congress Catalogue Card Number 74 79009

fi

All rights strictly reserved.

No part of this book may be reproduced
in any form without theJ'ermission in
writing of the publisher.'

2. The structure of hairs 5

Cross-sectional appearance
The medulla 5
The cortex 8
The cuticle 8
Hair pigmentation 10

Designed by Arthur Stohs

Text set in Monotype Times Roman by

Monotrade Pty Ltd
Melbourne, Victoria

Printed by Shanghai Printing Press

Hong Kong

4
5

SECTION B
1. Methods of studying hair structure 13
The wbole mount 13
Obtaining the Cross-section 13
Obtaining cuticular scale casts 14
Photographing hair structures 16

Ltd.~

I
I

I
i

2. A system for bair identification 17

Producing the system 17
Using the system 18

SECTION C
1. A guide to hai, structure for the indigenous and introduced
mammals of Victoria 19
2. A grouping of Victorian mammals hased initially on the crosssectional appearance of primary guard hairs 20
3. Photographs and descriptions of hairs for each species involved 22
Bibliography 173
Index to Species 175

SECTION A

1 Introduction

GENERAL

There are many situations where mammal identification

is most conveniently based on a consideration of hair
structure. The identification of ingested hairs is a
particularly useful aid in the determination of diet for
wild carnivorous animals, as most meals of mammal
origin would include at least a small quantity of hair.
Bones and other indigestible items are often avoided.
Ingested bones, with the possible exception of those
from very small prey species, are generally fragmented
and of little diagnostic value. By contrast, hairs suffer
much less damage during the digestive processes, and
even hairs recovered from faeces retain many of the
features useful for identification. Indeed, faecal analysis
is now a widely used teChnique in food habits and dietary
studies.
The hair from a number of mammals is commercially
utilized and a knowledge of hair structure is necessary in
certain sections of the textile and allied industries. Hair
identification may also play some part in the field of
forensic science and the policing of wildlife laws. In
mammal survey investigations, hair traces on twigs,
fences, etc. may weI! provide the only clue to the presence
of a particular mammal in the study area.
Although the history of hair structure studies related
to mammal identification may be traced back to the
late nineteenth century, the first significant contribution
in this field was the work of Hausman (1920,1924,1930)
in America. Using written descriptions and somewhat
simplified drawings, Hausman presented data on the
configuration of cuticular scales and medullae for the
different hairs of 166 fur-bearing mammals. However,
he made no attempt to provide any key as an identification aid and relied solely on a comparison between
known and unknown samples.

In the following decade, Mathiak (1938) produced

a key to the identification of hairs of the mammals of
southern Michigan. The key was based partly on an
appreciation of cross-sectional appearance and medulla
arrangement, but also relied heavily on measurements
of hair diameter and hair length. The major shor1eoming
of Mathiak's key stems from his heavy dependence on
these hair measurements as diagnostic aids. In the same
year, Williams (1938) produced a key to the identification
of hairs of moles and shrews.
At about this time, technological advances in the
textile industry prompted detailed hair structure studies
by the Wool Industries Research Association at Leeds,
England. Wildman (1940) published a booklet on animal
fibres of industrial importance and discussed their origin
and identification. This work was greatly enlarged and
repUblished in 1954. This publication has generally been
accepted as the definitive work on hair structure.
Understandably, Wildman's investigations centred about
a detailed description of fibres of commercial importance,
and no attempt was made to produce a key or system of
identification for unknown hairs. A further account of
this work was giv<ln by Appleyard (1960) in his Guide to
the identification of animal fibres.
A descriptive key to the identification of hair of
Californian mammals .was published by Mayer (1952).
The entire key was based on a consideration of dorsal
guard hairs which had been taken from one small area on
the pelage. In all, some 392 species and subspecies were
considered. Although this key was well devised, it relied
heavily on the value of hair length and general profile as
a diagnostic aid and ignored the value of crosssectional
shape and scale pattern.
Studies on hair structure of Tasmanian marsupials
and monotremes by Lyne and McMahon (1951) suggested

INTRODUCTION

THE IDENTIFICATION OF MAMMALIAN HAIR

that a detailed microscopic examination of hairs of these

mammals would be of some taxonomic value. In this
preliminary investigation they made no attempt to
produce a key to the hair of the spe<;ies involved. A
detailed study, dealing entirely with the hair of the
Chiroptera, was reported by Benedict (1957). Her work
involved a description of hair structure for 16 families
and also included a key to the dorsal hair of the Chiroptera, In more recent years studies On the diagnostIc value
of hairs have been reported by Day (1966) and Adorjan
and Kolenosky (1969).
These and many other studies have contributed geeatly
to our knowledge of mammalian hair and have de";'onstrated the use of hair structure as a diagnostic aid.
Despite this, there stil! remain a number of problems
associated with the identification of unknown hairs.
. These inolude the following:
1. the considerable variety of hair types encountered,
even on an indIvidual mammal;
2. variations in hair structure along the length of an
individual hair;
3. an appreciable degree of inter-species overlap, in
particular characteristics, limiting the usefulness of
some structure;,s as diagnostic aids, and
4. slow methods of study for some aspects of hair
structure which are used as 'diagnostic aids.

"

It is the aim of this book to present a particular

approach to hair identification, which attempts to minimize these problems. The major problem of inter-species
overlap is reduced by the use of a photographic reference
system. Although many of the mammals represented in
this photographic system (Section C) are endemic to
Australia, the general principles and methods outlined
could be expected to apply to any situation demanding a
rapid system for the routine identification of large
numbers of hair samples. The book has been primarily
designed with the needs of the food-habits investigator
in mind.

systematic significance. However, for the purposes of

identifying unknown hair samples, vibrissae are of no
value as their basic structure is very similar for all
mammals.
Bristle hairs On some mammals, such as certain
breeds of the domestic pig, the whole pelage is composed
of rather stout and rigid hairs which are of uniform
diameter along most of their length. Such hairs usually
have a very narrOw medulla, Or no medulla at all (see
page 36), are oval to circular in cross-section, and
commonly have a flagged tip. Although Noback (1951)
and others consider the term "'bristle to be synonymous
with protective hair, primary hair and overhair, it is
useful for our purposes to recognize bristles as being
different from the coarse hairs mentioned below.
H

Over/wirs A close inspection of most mammal pelages

will show that some sparsely scattered hairs are distinctly
longer than those comprising the bulk of the pelage.
These are termed overhairs. Usually they are circular in
cross-section and have little diagnostic value. However,
in those cases where their cross-SL"Ctional shape is not
circular, they may be useful in identification. As a rule,
overhairs are mOre densely pigmented than the other
hair types immediately surrounding them.
Guard hairs These are the larger or coarser of those
hairs forming the main pelage and include a type often
described as shield hairs. In shield hairs the distal part'
is noticeably wider and flattened, forming a shield.
Guard hairs lacking a shield are of uniform diameter
along most of their length, tapering only towards the
tip.
The largest of the guard hairs, conveniently termed the
primary guard hairs,. are of paramount importance in
hair identification, for it is these hairs which generally
exhibit the most diagnostically useful features. There is
usually a variety of sizes for guard hairs in anyone
pelage, ranging down until they are indistinguishable
from the underhairs (see below).
Underhairs These are shorter and finer than the guard

TYPES OF HAIR

For the purposes of determining the identity of a particular hair sample it is useful to recognize the following
"types" of hair:
Vibrissae These are large, stiff hairs that are primarily
sensory in function. They are variously referred to as
whiskers, sensory hairs~ tactile hairs and sinus hairs.
Generally speaking, the vibrissae are widest in their
proximal half and taper to a long tip. Lyne (1959) has
noticed that, in the Marsupialia, vibrissae have some

hairs on any given area of the pelage and are commonly

wavy. With few exceptions, they retain the same thickness
along their whole length (except the tip). Like the guard
hairs, they range in size down to extremely fine hair
which is sometimes called veUus or lanugo (Danforth,
1939). Underhairs are generally of little diagnostic value.
h should be pointed out that some mammals have a
much wider range of hair types than do others. For
instance; in the main pelage, rodents have more types
of hair than do the more common breeds of sheep.
Furthermore, the relative abundance of overhairs,

FIgure J A tuftofhairfrom the brown rat (Rallusnorveg<'cus)

showing an overhair (A), guard hairs (B), and
underhairs (C).

Occasionally, the underhairs exhibit particular features

of profile which are useful in confirming an identification.
C is rather an uncommon profile, representatiVe of an
underhair from the mouse (Mus musculus). Note the
constrictions at several points along the length of the
hair. At each constriction, the fibre changes direction,
giving an overall "zig-zag" pattern. In D, one of these
has been enlarged to show its form more closely. Constrictions vary greatly in their acuteness. Some are
narrow in both the horizontal and vertical plane whilst
others are noticeably more narrow in one plane than
in the other.
Many guard hairs are wavy in outline, and usually
the wave plane corresponds to that axis in which the
hair is of minimum diameter. Exceptions to this are
guard hairs from deer and goats, where the wave plane
is at right angles to that axis in which the hair is of
minimum diameter. Guard hairs with a prominent
shield are usually straight and stiffened in the region of
the shield.
Another useful diagnostic criterion, with respect to
the hair profile, is the length and width of the shield in
relation to that of the remaining portion of the shaft
Some guard hairs (e.g. in the house mouse, Mus
musculus) have a distinct long; tumnal channel in the
shield region (see page 15BA) and such hairs are arranged
so that this channel is always on the outer surface. The
hairs are commonly curved and the channel is invariably
on the convex surface of the curve.

guard hairs and underhairs in the pelage of any mammal

can change with time, the changes being associated with
age, moulting, nutrition, etc.
Most classifications of hair types are subjective in
their approach and, therefore, cannot hope to cover all
hairs on a given pelage. Irrespe<;tive of the system of
classification. there are animals where a continuous
gradation exists from one hair "type" to another.

THE HAlR PROFILE

The classification of hair types, as outlined above,

relies heavily on an appreciation of the profile or general
outline of a hair. A study of general profile may also be
of great value in attempting to identify an unknown
hair sample.
Some examples of profile are shown in Figure 2.
A is a shield type guard hair and is commOn to a great
variety of mammal species. B has a rather unusual
profile and is characteristic of the platypus (OrnUhorhynchus anatinus). Note the distinct constriction
immediately before the shield and, also, the round tip.

Figure 2 Some examples of hair profiles. A and B are guard

hairs whilst C is an underhair. D represents a
magnIfied view of a constriction in an underhair.

~""~~<-<~~-==:::::::--------------~-----------------------

THE IDENTIFICATION OF MAMMALIAN HAIR

2 The Structure
of Hairs
,,

A.

_A

fo.-

Scah!! Patterns

/1I

Djfferent

,I

A_

Skin
Cross~sectlons

A-A --_.

Growth Stages

Figure 3 Five stages iI~ the development of a primary guard

hair in the brown rat (Rattus narvcgicus),

ASPECTS OF HAIR GROWTH

Very often, hairs which have to be identified originate

from immature animals, It is~ therefore? essential to have
some understanding of hajr groMh in relation to pelage
changes.
The development of follicles and the early stages of
hair growth have been the subjects of numerous papers.
A detailed account is given by Lyne (1966).
These initial stages need concern us little, but the
sequence of appearance of various hair types in the pelage
is of particular importance. Quite obviously, this sequence
will vary considerably between species and may even vary
between< individuals of the one species.
As an example, however, we describe the sequence
of appearance of various hair types in the brown rat
(Rallus norvegicus). When the rat is about 3 weeks old
it is possible to recognize in the pelage, small overhairs,
small guard hairs and underhairs. At about 9 weeks of
age, larger guard hairs can be found. The largest guard
hairs (termed primary guard hairs in this book) are not
found fully developed until the rat is approximatly 3
months old.

With respect to hair identification, the implication of

this sequence of appearance is clear. In young rats the
most useful or diagnostic hair type (Le. primary guard
hair) may not be present and problems of identification
may be greatly increased.
It is important to realize that, once a hair has protruded
above the skin, it is no longer composed of living cells
and its detailed morphology will not then change.
Hence, the profile depends entirely on the activities of the
hair follicle. Some results of this are shown in Figure 3,
which depicts the growth of a primary guard hair
.
(shield type).
An important diagnostic character is that the crosssectional appearance of the hair at the widest point,
despite changes in hair length, remains unaltered during
growth. Hence it is always advisable to examine the
cross-sectional shape at the widest point along the hair
when inspecting unknown samples. It should also be
mentioned that changes in certain other structuml
features occur during hair gro'wth. These variations are
discussed later in the text.

The internal structure of hairs is probably best appreciated from a consideration of the cross-sectional appearance
in Figure 4. Typical hairs are made up of three layers of
keratin material: the central core or medulla, a layer of
cortex surrounding the medulla, and an outermost layer
called the cuticle or scale layer. All three layers are
composed of dead cells and all show their cellular nature
under detailed inspection.
The shape, arrangement and relative size of all thrce
layers is of great importance in hair identification. It
should be pointed out that some hairs are non-medullat-'
ed. Examples include the wool from some breeds of
sheep and hair from small bats. In some species, (e.g.
wombat, Vombatus ursin"s), the medulla may be found
in the distal half of the hair only.

CROSS~SECTIONAL

APPEARANCE

In addition to struCtural differences involving the cortex,

medulla or cuticle, there are quite marked differences in

the cross-sectional shape of different hairs. In general,

the most diagnostically useful cross-sectional shapes are
found at the widest point on the primary guard hair.
Cross-sections taken at other points along the hair have
a useful supplementary value in hair identification. It
should be realized, however, that the shape may change
along the length of the hair. Mathiak (1938) has demonstrated this quile well by producing serial cross-sections
which cover the variation in shape along an individual
hair.
The shape of the hair cross-section is undoubtedly
the most important single criterion used in the hair
identification system outlined in Section C It is also of
particular use in the preliminary grouping of mammal
species for a system of rapid identification of hairs (see
page 20) and is often sufficient to identify unknown hairs
without recourse to other hair structures.
The most common cross~sectional shapes encountered
are shown in Figure 5. Accurate measurements of hair
width for both the major and minor axes can only be
properly achieved by reference to cross-sections.
THE MEDULLA

Cuticle ---.<'J

Medulla ---/tI-Cortex----'i~

Figure 4 Basic hair structure.

The medulla is comprised of shrunken ceUs which may

or may not contain pigment. Spaces between these ceUs
are usually filled with air (or other gas) and appear as
obvious dark areas under the microscope. These dark
spaces can obscure the actual structure of the medulla,
and to ~'Ilable the structures to be seen clearly it may be
necessary to "infiltrate~' the hair, replacing the air or gas
in the medulla with a suitable medium (see page 7). In
most hairs, however, sufficient detail can be obtained
without recourse to this technique.
The general appeamnce of a medulla varies enormously
depending on the species from which the hair originated,

THE STRUCTURE OF HAIRS

THE STRUCTURE OF HAIRS

Circular
Medium size
Medulla

Circular
Large Medulla

o
Oval
Large Medulla

Oval
Medium size
Medulla

Oval
Medulla Absent

Eye-shaped

Narrow
medulla
lattice

Wide
medulla
lattice

Narrow
Aeriform
lattice

D
Wide
Aeriform
lattice

E
Simple

Interrupted

,
Oblong
Large Medulla

Oblong
Medium size
Medulla

Cigar-shaped

Concavo~convex

Concavo -convex
Large Medu lIa

Divided
Medulla

Bilobed Medu lIa

Concavo-convex

Reniform

~
Dumb-bell
Shaped

I
t
.

G
Fragmental

Uniserial
ladder

Multiserial
ladder

Globular

Stellate

Intruding

Figure 5 Some shapes of cross:-sections at the widest part of

the primary guard hair.

Figure 6 Medulla types.

the type of hair, and the point along the hair at which the
medulla is vievied. Differences in appearance also occur
between normal and "infiltrated" medullae.

(1954).

The Classification oj Medullae

I.

By reference to the general shape and arrangement of

both air spaces and medullary material, it is possible to
recognize and define four major structural groups of hair
medullae: unbroken, broken, ladder and miscellaneous.
Each of these can be further subdivided into a number of
medulla types. A total of 12 distinct types (Figure 6) is
defined below. Note that, for each type, the diagram
indicates the appearance of both a normal and "inftltrated" medulla. The classification used hereunder has

These consist of a continuous central core of a regular

or irregnlar diameter which can range from narrow
to very wide. Several typ<?S are recognized:
(I) Medulla Iotrice This is probably the most Common
type of unbroken medulla. The shrunken medullary cells
form a network or lattice and enclose air spaces of various
shapes. For convenience we may speak of a narrow
medulla lattice (medulla less than 1 of hair width)
(Figure 6A) or a wide medulla lattice (Figure 6B).

been adapted and supplemented from that of Wildman

Unbroken medullae

(ii) Aeriform lattice This relers 10 the type of medulla

where air spaces appear as a network or lattice, enclosing
aggregations of shrunken medullary cells. Many rodents
possess hairs with the aeriform lattice type of medulla
and the term "rodent base" has been used to describe
this particular arrangement (Mathiak 1938). Hausman
(1920) and otbers have classified this as a "compound
medulla". Again, for convenience, the aeriform lattice
may be further classified as narrow (Fignre 6C) or wide
(Figure 60). In many mammals, the lattice type medullae
are found only in the wider parts of hairs.
(iii) Simple This type has no obvious medullary
structure and the medulla area may be relatively narrow,
or wide (Figure 6E). The width may vary along the

length of an individual hair. Quite often hairs with

a simple medulla are found in association with hairs
having a lattice type of medulla. It should be pointed
out that damaged hairs (e.g. from faecal samples) often
show an apparent simple medulla due to the destruction
of medullary networks by the digestive processes.
2.

Broken medullae

Here, the medulla is not continuous along the length

of the hair, but is interrupted by sections of cortical
materiaL Two types may be recognized:
(I) Interrupted The medullary column is interrupted
along its length by short sections of cortical material
(Fignre 6F).

THE IDENTIFICATION OF MAMMALIAN HAIR

(ii) Fragmented In this type the medullary column is

interrupted by long sections of cortical tissue, The
isolated medulla sections show little differentiation and
their structure could be termed simple (Figure 6G).

may be an important diagnostic criterion, Very often too,

the ratio of cortex width to medulla width (shown in
cross-section) can be of use in the identification process.

3, Ladder medullae

THE CUTICLE

This classification includes those medullae having one'

or more rows of air spaces. Two distinct types can be
defined:
.
(i) Uniserialladder The medulla has a single row of air
spaces and these can be angular, rounded, flattened or
cup-shaped (Figure 6H). This type is .usually found in
underhairs and the proximal part of Some oU,er hair
types,
(li) Multiserial ladder The medulla has tWo or more
distinct rows of mostly uniform air spaces (Figure 61),
Such a med,ulla is characteristic of lagomorph hairs,

The cuticle or outer layer of a hair consists' of a single

layer of generally transparent, overlapping scales. In
rare cases, the scales may be pigmented (e.g. some
Chiroptera-Benedict 1957), Along most of the hair
length, scales are flattened against the body of the hair,
with their free edges pointing towards the hair tip. At
the base of the hair, however, scales may be more "open"
in arrangement and a cross-section in this region shows
the cuticle to have a rippled or jagged outline, with each
ripple corresponding to an individual seale, In this sense,
the surface of the hair may be likened to that of a pine
cone, where seed receptacles near the base of the cone
are partly opened and those towards the tip are tightly
closed,
An unusual arrangement of scales can be observed
on hairs originating from the foot pad of some lagomorphs. Here, the scales are positioned at a considerable
angle to the long axis of the hair and are tightly packed.
This has the effect of producing an extremely thick
cuticular layer, the advantage of which is obvious in
such a situation,
The size and shape of scales, and their pattern of
arrangement around the hair, are useful criteria for
identification purposes, In relation to scale size, a
measurement termed the scale index has been used by
some workers (Hausman 1930, Mayer 1952). This can
be defined as the ratio of the free proximodistal length
of a scale to the diameter of the hair shaft. This measurement is of doubtful value as a diagnostic aid. Along
quite short distances of an individual hair, scale sizes
may vary considerably, with a resultant change in the
scale index. Furthermore, such a measurement conveys
no information on the actual scale shape (i,e. a short
and wide scale on a thin hair might have the same index
as a long and narrow scale on a wide hair), although
Khemelevskaya (1965) has suggested that the index
might be of some u.e if a measurement of scale width
was included.
A manual for hair identification, based almost entirely
on photographs of scale shapes and arf'dngements, has
been published by Adorjan and Kolenosky (1969), and
many other workers depend heavily on aspects of the
cuticle in attempting to identify unknown hairs.

4,

Miscellaneous medullae

These are generally uncommon and represented in the

main pelage of only a small number of mammals or
restricted to hairs taken from extremities (e.g, tail), The
following types may be recognized:
(i) Globular This type has an aggregation of globular
air spaces (Figure 6J). It may be found in SOme hairs
from the platypus (Ornillwrhynchus analiflus), wombat
(Vambatus ursinus) and fur seal (Arctacephalus doriferus).
(ii) Stellate Here, an essentially "simple" medulla has
finger-like projectiorts radiating out into the cortex
(Figure 6K). This type is often found in the tail hairs of
larger mammals, and is usually best appreciated in
cross-sections.
(iii) intruding This type has narrow and irregularly
placed air spaces projecting into the cortex. The spaces
may project in several directions and are not necessarily
found in the centre of the hair (Figure 6L),
THE CORTEX

Like the medulla, the cortex of the hair is composed

of dead cells, but the individual cells are usually not
visible under the light microscope since they are packed
into a rigid and almost homogeneous hyaline mass
(Hausman 1932), Due to a high degree of cornification,
the cortex has a low reftactive index and, in the absence
of pigment, is translucent. It is sometimes possible (e,g,
by retting wool fibres) to discern outlines of individual
cortical cells, and these are generally polygonal in shape.
However, few data are available on the way in which
cortical cells are arranged in various types of hair,
.Because the cortex lacks discernible structure, under
the light microscope, it is of little value for identification
purposes, However, in those hairs where the cortex is
pigmented, the size and arrangement of pigment granules

The classification of cuticular scales and scale patte,.s

The scheme of classification shown hereunder is based on

the following main features:
I, Terms which describe the form of the scale margin,

THE STRUCTURE OFHAIRS

FORM OF SCALE MARGINS

A. Smooth

B. Crenate

C. Rippled . D. Scalloped

DISTANCE BETWEEN

g;q

E. Dentate

SC ALE MARGINS

F, Distant

G, Near

SCALE

H. Close

PATTE RU
r

~'2:
,
I. Simple
Coronal

O. Regular
Wave

J, Diamond

K, Narrow

Petal

Diamond Petal

p, Irregular
Wave

Q, Single

Chevron

L Broad
Petal

M. Regular
Mosaic

R. Double

N. Flattened
Irregular
Mosaic

T, Transitional

Chevron

Figure 7 Various. shapes and arrangements of cuticular

scales.

2,

Terms which describe the distance between the

external margins of scales.
3. Terms descriptive of the general scale patterns.
The scheme as outlined, is generally similar to that of
Wildman (1954), although some additions and modifications have been made, All scale patterns described have
been based on an appreciation of scale casts rather than
rolled impressions (see page 15 for an explanation of
these terms),
Before considering detailed aspects of scale classification, it is useful to recognize that there are 'two broad

groups of scales; the coronal group and the imbricate

group, A coronal scale completely encircles the hair
shaft, whilst an imbricate scale extends only part of the
way round the shaft circumference, It follows that
several imbricate scales may be required in order to
span the circumference of the hair at any point along its
length. An example of each type described is shown in
Figure 7, (I--coronal, J-T-imbricate),
L

Scale margins

The scale margin may be defined as the free distal

edge of an individual scale,

THE IDENTIFICATION OF MAMMALIAN HAIR

Smooth margins are without indentations and appear
as a smooth line (Figure 7A).
Crenate margins have shallow but relatively pointed
indentations (Figure 7 B).
Rippled margins have deeper indentations but the
profile is generally rounded (Figure 7 C).
Scal/oped margins consist of a series of curves with
generally rounded peaks and pointed troughs (Figure
70).
Denlate margins have large tooth-like projections and

are found only on coronal scales (Figure 7 E).

Distance between margins (Figure 7 F-H).
In describing the arrangement of cuticular scales, it may
be desirable to indicate whether the free edge of one
scale and that of adjacent scales above and below are
close together or well separated. For convenience we
may speak' of distant, near and close scale margins.
these being purely arbitrary divisions. The distant type
are those where the visible width of the scale is not more
than three times the visible length (i.e. the distance
between one free distal scale margin and the next). In
near types, the ratio of width to length might be Irom 3 to
about 8, and in c/ose types it is greater than about 8.
Scales in the proximal half of a hair are commonly
longer (i.e. more distant than those in the distal half).
2.

3. Scale patterns
The majority of harfs used for identification purposeS
have cuticular scales of the imbricate type, and these are
arranged in a variety of patterns, around the perimeter
of the hair. Patterns may change markedly along the
length of the hair with a corresponding change in the
appearance of individual scales.
For anyone hair, an examination of the cuticle may
reveal one or more of the following scale patterns:
I. Petal types Wildman (1954) described the general
appearance of these as being similar to the patterns
formed by a series of overlapping flower petals. Very
often, the individual scales may appear to be diarnondshaped. In these cases, the pattern can be termed diamond
petal (Figure 7. J-K). The seales forming this partkular
pattern can range from being wide and short to being
narrow and long. Another type of petal pattern, termed
the broad petal (Figure 7 L), can be recognized, where the
individual scales have rounded distal edges and are often
of irregular sizes.
2. Mosaic types These patterns have an overall
angular appearance. The term mosaic, as applied to the
appearance of the pattern, is self explanatory. Sometimes,
the width of individual scales is similar to their length
(i.e. visible length and width) and we may speak of

THE STRUCTURE OF HAIRS

regular mosaic patterns (Figure 7 M). Hence, irregular

mosaic patterns occur when individual scales forming
the pattern are of dissimilar sizes and shapes. It is also
useful to recognize another common type of mosaic
pattern, namely the flattened mosaic. The irregular
mosaic pattern shOwn in Figure 7 N is also a flattened
mosaic. Here, individual scales appear to be much wider
than long (near), but still form an angular mosaic
arrangement.

a variety of colour hues, with black, brown and yellow

being the most common or basic colours (Fitzpatrick
ot al. 1958).
Pigment type
It is possible to describe the presence of pigment
in a hair as either diffuse, granular or aggregate; these
terms being self-explanatory. Granular or aggregate
pigment may be arranged in discrete and well defined
areas Or in a streaked pattern. Individual granules may be
large or small. Figure 8 gives an example of large
granules of pigment in the cortex.
The use of pigmentation as a diagnostic aid in hair
structure is subject to many limitations. For anyone
mammal, there may be a noticeable variation in colour
between hairs from different parts of the body. Again,
the distribution of pigment may change considerably
along the length of the hair. Colour mutations and fading
are some other factors which limit the use of pigment
in identification of unknown hairs.
The pelage of certain mammals will fluoresce under
ultra-violet light, and Latham (1953) has used this
char~cteristic to separate pelts of the least weasel
(Mustela rixosa) fwm those o.f two other species in the
genus. However, this phenomenon appears to be quite
restricted and only oflimited application in identification
of mammalian hair.
'

3. Waved types On some hairs, the free distal edges

of the scales are presented in such a fashion as to appear
like a series of waves. For all imbricate scales, the
waves are not continuous but are interrupted by virtue
of scale overlap. A true continuous wave can only be
observed when rolled impressions (see page 15) are made
from hairs with coronal scales. Waves may be either
regular (Figure 7 0) or irregular (Figure 7 P) in appearance and, depending on their amplituc\e, may be desscribed as shallow, medium or deep. In cases of irregular
waves, the amplitude of successive waves changes
noticeably. Other types of waves have been identified
and described by Wildman (1954). These include the
single and double chevron (Figure 7 Q-R), and the
streaked ware (Figure 7 S). In the single chevrqn, either
the crests or troughs are pointed. whilst in the dou hie
chevron, both crests and troughs are pointed and the
Waves tend to have a somewhat larger amplitude. The
streaked wave pattern has alternating series of deep and
shallower waves and the change in wave form corresponds
to a change in the contour of the hair circumference.
The streaked wave pattern can often be observed on the
outer surface of those guard hairs having a longitudinal
channel (see page 1670).

Figure 8 Cross-sectional appearance of hairs of the euro.

(Macropus robustus) showing prominent pigment
granules.

4. Transitional patterns (Figure 7 T). By reference to

Figure 3 (page 4), it will be noticed that the scale
pattern changes markedly as one progresses from the tip
to the root of an individual hair. At those points where
one type of scale pattern changes into another, the
pattern is often termed transitional. Some actual examples
are shown on pages 67 and 69. Note that the transition
from one pattern to another can take place over a very
short distance along the hair.

Pigment distribution

Pigment may be present in the cortex or medulla

or both regions of a hair and there are isolated occasions
when cuticular scales may be pigmented (Benedict 1957).
An important aspect is the way in which the pigment is
distributed along the length of an individual hair. Many
mammalian hairs present a banded appearance and both
Stains (1958) and Mayer (1952) have utilized aspects of
colour banding in their keys to the identification of
hairs. Again, the location of pigments in hair crosssections may be of importance, and Dreyer (1966) has
produced a detailed classification of pigment distribution
based on an appreciation of cross-sections.

HAIR PIGMENTATION

Although not of primary significance as a diagnostic

criterion, hair pigment and its distribution can often be
of use to confirm an identification made on the basis of
other hair characteristics. In isolated cases, the particular
. appearance of hair pigment may be quite characteristic
of a certain species and hance, be of great value in
identification (Figure 8).

Pigment colour

Two pigment types are known to occur in mammalian

hair: eumelanin and pheomelanin. Eumelanin results
in brown or black hair, whilst pheomelanin gives a
yellow or reddish colour. Despite this, hairs may display

t,,'

10

"

[I

11

----------,----------------

-~-~~~~
...

SECTION B

1 Methods
of Studying
Hair Structure

THE WHOLE MO{;NT

When hairs are mounted on a microscope slide, it is

generally possible to observe not only the general profile
of the hair but also the appearance of the medulla (if
present) and the distribution of pigment.
If the hair sample is taken from a study skin and is to
be used as a reference, several precautions are necessary.
First, a full range of bail's (i.e. a tuft of hairs) should be
selected from each of several parts of the body and
removed from the pelt in such a manner as to ensure that
the maximum accessible length of each hair has been
obtained. This can be achieved by using a very sharp
razor blade and cutting flush to the skin surface.
The hairs must be cleaned, and 70 % alcohol or an
alcohol/ether mixture (equal proportions) is useful for
this purpose. Dried hairs can then be mounted .in a
suitable medium on a clean glass slide. Several mounting
media are available and the merits of each have been
outlined by Wildman (1954). For reference slides, a
permanent mountant such as DePeX (Gurr) is the most
useful. When mounting the hair tuft, it is necessary to
ensure that individual hairs are well separated so that
a "hair forest" situation is avoided. Depending on the
length of individual hairs, it may be necessary to use very
long coverslips or to cut the hairs into two or more
sections and mount each section in order. This technique
of cutting hairs is also used to observe infiltrated medullae,
as the mounting medium will. infiltrate the medullary
spaces for a short distance from the cut end.
F or routine identification work (Le. preparing mounts
of unknown hairs) it is generally more practicable to usc
temporary mounting procedures. This allows for the
re-use of glass slides and, also, is considerably faster than
using permanent mounting. If possible, a tuft containing
the full range of hairs should be selected, cleaned and

mounted. Paraffin oil is a most convenient medium for

temporary mounts. An important advantage of the
temporary mount is that hairs can subsequently be
removed from the slide and used for othar purposes
(e.g. cross-sectioning). This is of particular value when
the unknown sample contains only a small number of
individual hairs.

OBTAINING THE CROSS-SECTION

A number of methods are available for obtaining the

hair cross~section and these vary from simple hand
sectioning techniques through to the use of speci~lIy
designed microtomes.
Where thin sections are required, it is usual to emp'loy
fibre microtomes. Several such instruments bave been
devised and they are widely used in the field of commercial
fibre microscopy. A brief description of these instruments
has been given by Wildman (1954) and Stoves (1957).
For the purposes of bair identification, very thin
sections are not required and simple methods of sectioning can be used. Hand sectioning has been successfully
employed by Mathiak (1938) and many others, but the
most efficient of these simpler techniques would appear
to be a modification of the plate method described by
Ford and Simmens (1959). All cross-sections photographed or described in this book were obtained by the
use of this technique,
The plate method
The apparatus consists of a stainless steel microscope
slide of approximate dimensions 76 X 25 X O 5 mm with
two to six holes, each of 08 mm diameter, drilled at
equal intervals along the centre as shown in Figure 9.
These holes are carefully contoured on the top and

13

METHODS OF STUDYING HAIR STRUCTURE

THE IDENTIFICATION OF MAMMALIAN HAIR

Figure 9 The plate method of cross-section;ng (see text).

bottom, to eliminate rough edges. The finished plate

must he kept highly p~lished and perfectly flat.
For sectioning, a loop of thin nylon thread (approx.
diam. 03 mm) is passed up through one of the holes
(Fig. 9A) and threads of cellulose acetate yarn (330
decitex) are inserted into the loop (Fig. 9B). These
threads are pulled a short distance down through the
hole with the aid of the nylon thread. The hair tuft for
identification, which should include a range of guard
and underhairs, is then inserted into the centre of the
yarn bundle protruding from the top of the plate (Fig.
9C). Hairs and packing material can then be pulled any
desired distance through the plate by means of the
nylon yarn.
The cross-sections are obtained by using a degreased
safety razor blade to cut the protruding hair bundle
flush to the plate on both sides. The blade is held at about
35 degrees to the plate (Fig, 9C), The cross-sections are
viewed in situ after placing paraffin oil and a coverslip
directly over the sectioning hole, Sections should be
viewed from that side of the plate upon which the second
cut is made (i.e, the underside),
In some samples, where large hairs have only a thin
or narrow cortex layer, the shape of the hair in the
cross-section may become distorted as a result of undue
packing pressure in the sectioning hole, Such distortion
can be overcome by using fewer hairs and dissolving

14

the packing material with a few drops of cellulose

acetate solution (2~A %in acetone) immediately prior to
cutting tbe section, This relieves undue pressure and the
dissolved yarn resets in a few second" thus preventing
any part of the bair bundle from falling out of the plate.
The plate method has a number of distinct advantages
over other sectioning methods, It is rapid, and the
experienced operator can produce cross-sections ready
for microscopic examination within two minutes, Very
short hairs (such as are often found on the head, reet and
ears) and damaged hairs, can be cross-sectioned with
relative ease, Because cross-sectional appearance often
varies along the length of an individual hair, longer
hairs can be preent into sections before they are inserted
in the plate, For a permanent record, plates can be
sealed and labelled. A particular advantage of the plate
method is that the plate can he re-used indefinitely simply
by pushing out the contents of the sectioning hole with
a mounted needle.

Procedures for studying scale patterns usually involve

the use of special media to obtain a cast or impression of
the actual hair surface, These are often produced directly
on to glass slides and viewed under the microscope,
Wildman (1954) differentiates between the terms "cast"
and "irnpressionH and restricts the use of the latter tenn
to those cases where the entire circumference of the hair
is represented, Hence a scale cast refers to the situation
where only part of the hair circumference is represented,
Impressions are obtained by carefully rolling the hair
over the surface of a suitable medium on a slide so that
the scale pattern around the full circumference of the
hair can he appreciated. Whilst this may be desirable, the
technique is rather time-consuming and of limited
application in routine hair identification work. Very
short hairs, damaged hairs and hairs which are extremely
flattened in cross-section can be accurately rolled only
with the greatest difficulty,
[t is more efficient, in hair identification work, to
employ the "scale cast" technique, Here, the hair is
placed on or pressed into a suitable medium which will
then reproduce the surface structure for that part of the
hair in contact with the medium. Various media are
available, including gelatin, celluloid, vinyl acetate
polymers and various specialized products such as
ethofoi! and permount (Wildman 1954).
For general work, the use of polyvinYl acetate (pva) or
vinyl acetate copolymers is recommended. These have
several advantages, They are inexpensive, easily obtained
and readily soluble in water, Moreover, they have
excellent casting properties and will faithfully reproduce
surface structures, A large range of such polymers is
available commercially and the particular choice is
hest left to the individual,

with a thin layer of diluted pva, using a fine flat brush.

Casts are not produced directly onto standard microscope slides because the thickness of the glass will not
allow viewing of inverted casts at high magnifications,
The pva has good levelling qualities and will produce a
film of fairly even thickness. A 50 % solution of pva in
distilled water is generally adequate, although more
concentrated solutions may be necessary for very coarse
hairs.
The selected hairs should be cleaned in an alcohol-ether
mixture and dried between good quality absorbent paper.
The individual hairs are conveniently handled with fine
forceps and a mounted needle. Several hairs should he
placed on the coverslip, some with tips overhanging and
others with the hair root or lower shaft region overhanging (Figure 10), All hairs should he gently pressed
into the medium so that adequate surface contact is
ensured,
After some 20 minutes at room temperature, hairs
can be removed from the coverslip. This is effected by
grasping the overhanging section of each hair with
forceps and carefully lifting it out of the medium. For
viewing the casts, the coverslip is inverted and placed on
a normal microscope slide. Such an arrangement will
show the scale pattern as it would normally appear on
the hair surface (i.e, convex view),
Waved or twisted hairs must he treated with special

A simple casting technique

A long glass coverslip (50X20xO,2 mm) is coated

OBTAINING CUTICULAR SCALE CASTS

As a rule, the surface structure or scale pattern of a

hair cannot be properly appreciated from the whole
mount preparation, Exceptions occur in hairs which
have little or no medulla and are either non-pigmented
or only slightly pigmented.

-.=
... ==============
Figure 10 Obtaining the scale cast (see text),

15

~-----~-~-----""----------------------'!lIP-------------------

THE IDENTIFICATION OF MAMMALIAN HAIR

care if useful casts are to be produced. It is generally

advisable to fasten both ends of such hairs with small
pieces of gummed paper so that they are correctly
positioned for embedding and are laid fiat along their
entire length.
In the case of very fine hairs (e.g. of some Chiroptera)
the strong adhesive property of pva prevents removal of
the hair from the coverslip. It is then advisable to use a
gelatin solution in place of pva. A coverslip is coated with
a thin film of gelatin solution (3 % in water)_ Hairs are
laid out in the manner described above and stripped from
the glass when the medium has dried sufficiently.
When studying scale casts it is important to view the
scale pattern at various points along the length of the
hair, and the points of transition between one type of
scale pattern and another. It must also be appreciated
that some hairs have recognisable "sides" (e.g. a COncave
and a convex surface) and that such hairs have a tendency
to consistently present only one "side" to the embedding
medium. Such a problem can be overcome by deliberate
. manipulation of hairs whilst they are being laid in the
embedding niedi urn.
Scale casts cannot be successfully stored for long
periods and it is therefore essential that photographic
records are kept for comparative work.
PliQTOGRAPHlNG HAIR STRUCTURES

It is not intended ifere to deal with the subject of

photomicrography in any detail but, rather, to mention
some of the general principles involved in producing
photographs for use in a hair identification system. The
reader is advised to consult more specialized texts for
information on photographic equipment, film types,
lighting etc. Some useful technical information on
photomicrography of hair is given by Appleyard (1954).
The photographs of hair structure in Section C of this
book were produced by the use of a 5" X 4" bellows
camera using cut films in a double-sided holder_ All
photographs of hair cross-sections, whole mounts and
scale casts repreSi'nt a total magnification of 468 (3).
Large format cameras allow for contact printing from
negatives and enlargements should seldom be necessary.
Nevertheless, modern film in 35 mm cameras should
produce negatives of a high standard, allowing for
considerable enlargement without loss of detail.
It is important that all photographs of hair structure
be produced at the one magnification. This allows for
direct comparison between photographs and is par,
ticularly useful when measuring the maximum diameter
of primary guard hairs from different species.
Although colour transparencies of hair structures
(particularly cross-sections) may be of use in the identification process, the assessment of colour is difficult and

16

intra-species variation in hair colour may be considerabJe.

For these reasons, colour photographs have not been

included in the photographic system outlined.
The photographs of hair profile shown in Section C
were taken after projecting the image of hairs on to a
white paper screen and carefully drawing the outline of
the hair. A standard 35 mm projector was used and the
hairs were held between two glass plates. The enlarged
drawings of hairs were subsequently placed over an
appropriate scale and photographed_

2 A System
for Hair
Identification

When hair identification is employed as a routine

technique, the system adopted must have the capacity to
screen a range of hair structures for all mammals likely
to be present in the particular situation or area under
study.
The identification can be effected by comparing the
unknown sample with the full range of known hair
samples. This direct comparison technique is, however,
tedious and slow and quite unsuitable for routine
analysis work.
One might consider the use of descriptive dichotomous
keys. Unfortunately, these have several shortcomings,
when applied to mammalian hairs~ Like the direct
comparison method, they are slow and often difficult to
interpret Their major drawback, however, stems from
the fact that they are unable to fully represent the intra
specific variation in hair structures that is characteristic
of most mammals.
One solution to these problem, is to photograph the
most diagnostic characteristics of hair structure for all
mammals concerned and use these photographs as a
reference system. Useful aspects of microscopic structure
for single hairs and groups of hairs can be represented and
appreciated directly from the photographs. This eliminates the tedious "procedure of direct comparison with
specimens of known and unknown hairs but, at the same
time, allows for greater representation of structure
variations than does a written key.
It must be remembered that no system of hair identification can be fully effective. Sometimes, hair structures
for closely related mammals appear identical in all
respects and no differentiation can be made. Again,
there are occasions when an unknown sample is devoid
of those hairs which exhibit the most diagnostic features
of structure (usually primary guard hairs). In these
cases, precise identification may not be possible.

PRODUCING THE SYSTEM

It is necessary, at the very begInning, to point out that

the construction of a photographic reference system and
its subsequent use~ requires some experience on the part
of the operator. It is only experience which allows one to
recognize the most diagnostic features of hair structure
for each species considered or to identify a particular type
of hair in an unknown sample (i.e. primary guard hair,
overhair etc.)~ The time taken to achieve this expertise,
will depend largely on the number of species which must
be represented in the system and the complexity of hair
structures encountered.
.
The first requirement for the construction of the
reference system is a set of accurately identified study
skins, representative of all animals likely to be encountered
in the identification work. The methods of preparing
study skins have been outlined by Dimpel (1971) and
many others, and are outside the scope of thip book.
Hair samples should normally be taken from the rump
area of each skin but, if hair structure differs markedly
from place to place on the species examined, it will be
necessary to include hair samples from other parts of the
body. Each sample must indude all hair types present in
the main pelage (i.e. overhairs, guard hairs and underhairs).
For most mammals, the following features of structure
should be carefully examined and photographed:
(a) the general profile of primary and smaller guard
hairs and an underhaif.
(b) the cross-sectional appearance of primary and
smaller guard hairs at their widest point (the cross'
sectional appearance of underhairs can also be
represented in this one photograph).
(c) the medulla arrangement for primary and smaller

17

,
THE IDENTIFICATION OF MAMMALIAN HAIR
guard hairs at several points along their length.
(d) the scale pattern at several points along the length
of a primary guard hair.
Where large numbers of photographs are required
to cover the four points mentioned above, it is usually
possible to ecOn lize by retaining only those particular
aspects of medulla arrangement, scale pattern, etc.
which appear to have the greatest diagnostic value.
The photographs of scale pattern, medulla arrangement
and cross-sectional shape for hairs from all the mammals
considered should be taken at the one standard magnification. Various magnifications can be chosen fur all
photographs of hair profile, but in each case a meaSurement scale should be included. The use of .,standard
magnifications will enable direct comparison between
photographs. and give some idea of relative sizes for
various hairs and their components.
In any reference system containing a relatively large
number of species (say in exceSS of 30), it should be
. possible to split up the reference system into several major
groups of mammals. This grouping can be carried out on
the basis of gross differences in the appearance of one or
more hair structures. In the case of the mammals which
the authors studied, the division was best effected on the
basis of differences in cross-sectional shape at the widest
point along the length of the primary guard hairs. In
other situations it may be more convenient to divide on
the basis of general '{profile of guard hsirs, medulla
arrangement or other hair characteristics.
Within each resulting group, it may be possible to
carry out further subdivision. For instance, in our
experience, some groups can be subdivided on the basis
of medulla appearance, presence of constrictions, presence
of divided medulla and other characteristics.
Such a division into groups and sub-groups substantiaJJy reduces the number of photographs which must be
scrutinized before an identification can be made. At the
same time, it is necessary to point out the danger of
excessive subgrouping. In effect this produces a situation
similar to that which results from attempting to use a full
descriptive key and introduces the difficulties inherent in
that system.

USING THE SYSTEM

For efficient identification it is best to have a tuft of

the unknown hairs. This should ensure that all desirable
hair types are present and, most importantly, will
facilitate the location of primary guard hairs. The
unknown sample may often contain hairs from more

18

than one species but, fortunately, hair tufts from each

mammal usually remain intact and differences in the
microscopic appearance of tufts will allow their separation.
When only small numbers of hairs are available or
when the sample has been extensively damaged, this
procedure of sorting out discrete groups or bundles of
hair will not be possible.
Whatever the situation, a range Of mixture of hairs
should be selected for closer examination. Whenever
possible, the selected hairs should include several of the
primary or larger guard hairs. For the sample so selected
it will then be necessary to produce whole mounts so that
aspects of medulla appearance, pigment distribution and
general profile can be observed. The most important
step, however, is the production of cross-sections. All
hairs in the selected sample (a hair tuft or mixture of hairs)
can be sectioned in the one operation. Where guard
hairs can be recognized in the sample, the cross-section
should be taken in such a way so as to ensure that these
hairs are sectioned at their widest point. When poor
samples are encountered, it may be necessary to repeat
the sectioning process until maximum representation of
all possible cross-sectional shapes present in the sample
can be achieved.
From an appreciation of cross-sectional appearance,
and to a lesser extent general profile, medulla, or other
hair characteristics, it should be possible to select quickly
the appropriate group or subgroup of mammals to which
the unknown hairs belong. Finally, however, it will be
necessary to compare aspects of the unknown sample
against a series of photomicrographs.
Scale patterns can very often be ignored in the identification process and are generally only used to confirm
identifications made on the basis of other criteria.
However, there will be some occasions when the scale
pattern is of primary diagnostic significance.
Notwithstanding the value of photographs, it is
always advisable to have, close at hand, a full set of
reference skins. Direct comparisons may be necessary
when the unknown hairs are suspected of having originated from extremities on the body (e.g. tail, foot, etc.).
These hairs will often be markedly different from those
on the main pelage and may not be represented in the
photographs.
The identification system, as outlined above, has been
used successfully by the authors in the examination of
several thousand hair samples from the stomach contents
of foxes (Vulpes vulpes L. J, dingoes (Canis jamiliaris
dingo) and feral domestic cats. (Coman 1972, 1973:
Coman and Brunner 1972). A large number of hairs
from faecal samples have also been successfully identified
using this system. (Brunner and Coman, unpublished
data).

SECTION C

1 A Guide to Hair Structure

for the Indigenous" and
Introduced t Mammals of Victoria

It is important to realize that the photographs in this

section represent only some of the multitude of hair

structures observed in the hair of anyone species.
Generally speaking, only the most diagnostic features
have been covered.
It should also be noted that with very few exceptions,
hairs from extremitjes) such as the taH. ear or mane, have
not been included in the system. Such hairs may be
encountered from time to time, especially in food-habits
investigations. Usually hairs from the extremities are
circular to oval in cross-section with a small medulla
and are often of a bristly nature.
For each species considered, a short commentary
is given and this attempts to point out particularly
useful features of structure. The maximum diameter of
guard hairs is also given and a scale, useful for measuring directly from photographs is located below.
Some arrangement of species into groups or sub-groups
(based on hair structures) has been possible. In each
group or sub-group, the species are arranged within
families or genera.
The following key has been used on 80me oflhe crosssection photographs:
G Primary Guard.hair
o Overhair
U Underhair
Micron

50

100

150

200

250

111111111111111111111111111
Scale for photomicrographs in this section. Each division equals 10 microns.

* Classification according to

Ryde (1970,

t Classification according to Walker (1964)

19

2 A Grouping
of Victorian Mammals
BASED INITIALLY ON THE CROSS-SECTIONAL
APPEARANCE OF PRIMARY GUARD HAIRS

'.

Family
. GROUP

Genus and Species

Hairs predominantly circular in cross section.

Dasyuridae

Phalangeridae

Dasyurus maculatus page 22

Dasyurus viverrz'n~s 24
Sminthopsis crassicaudata 26
Acrobates pygmaeus 28
Cercartetus concinnus 30
Cercartetus nallus 32
Burramys parvus 34
i;{'~

Hairs predominantly oval in cross-section.

Sub-group (a) Hairs with medulla much reduced or
absent.
Sus serofa 36
Suidae
Vambatus ursinus 38
Phascolomidae
Homo
sapiens 40
Hominidae
*Ovis
aries
42
Bovidae
Nyctophi!us geoffroyi 44
Vespertilionidae
Miniopterus schreibersii 46
J::-ptesicus pumilis 48
Chalinolobus gouldii 50
Myotis adversus 52
Sub-group (b) Hairs with very large medulla.
Dama dama 54
Cervidae
GROUP 2

* Axis porcinus 56
Sub-group (0) Hairs up to 451' (max. diam.) and with
distinct constriction before shield.
Dasyuridae
Antechinus j/avipes 58
Antechinus stuartii 60
Antecninus swainsonii 62
Anteehinus minimus 64
Sminthopsis murina 66
Sminthopsis leueapus 68

20

Family

Genus and Speceis

Sub-group (d) Hairs larger than 40/-, (max. diam.) and

without constrictions.
Macropodidae
Maeropus glgomeus 70
Maeropus fuliginosus me/anal's 72
Maeropus robustus 74
Macropus rufogrL,eus 76
Megalela rufa 78
Wallahla bieolor 80
Thylogale billardierii 82
Petrogale penicillata 84
Potorous tridactylus 86
Canidae
Canis familiaris 88
Vulpes vulpes 90
Equidae
'Equus wbollus 92
Felis eatus 94
Felidae
Bovidae
Bas taurus 96

3 Hairs with eye or lemon-shaped crosssections.

Phalangeridae
Trichosurus Quipecula 98
Trichosurus caninus 100
Petauridae
Pseudocheirus peregrinus 102
Pe/aurus breviceps 104
Pe/aurus nor/oleensis 106
Petaurus aus/ralis to8
Schomobates volans 110
Gymnobe/ideus leadbeateri 112
Phascolarctidae
Phascolaretos emereus 114
GROUP

GROUP

Genus and Species

Family

Sub-group (b). Medulla present.

'Ceraus un/color 118
Cervidae
'Cercus e/aphus 120
Mustelidae
Mustela pu/orius 122
Phascogale tapoatafa 124
Dasyuridae
Hydromys ehrysogaster 126
Muridae
Notomys miteheliii 128
Pseudomys novaehollandiae 130
Pseudomys a/boeinereus 132
Pseudomys fumeus 134
GROUP

in

5 Hairs predominantly flattened or cigar-shaped

cross~section,

Otariidae
Arctocephalus doriferus 136
Ornithorhynchidae Ornithorhynchus anat/nus 138
6 Hairs predominantly renifonn or cone"VOconvex in cross~section.
Sub-group (a) Hairs with divided medulla.
Peramelidae
Isoodon obesulus 140
Perameles nasuta 142
. Perameles gunnii 144
GROUP

Family

Genus and Species.

Hairs with bilobed or large medulla.

Rattus rattus 146
Rattus norvegicus 148
Rattus fuscipes 150
Rallus lutreolus 152
Pseudomys shortridgei 154
Mastacornys fuscus 156
'Mus musculus 158
'Cavia parcellus 160
Caviidae
Sub-group (c) Medulla much reduced or absent.
Pteropus scapulatus 162
Pteropodidae
Pter!'pus palioeephalus 164
Sub-group (b)
Muridae

GROUP 7

Hairs predominantly dumb-bell-shaped in

cross-section.

Bovidae
Leporidae

Capra hircus 166

Oryc/olagus cuniculus 168
Lepus europaeus 170

4 Hairs with predominantly oblong cross-

sections.

Sub-group (a) Medulla absent.

Taehyg10ssidae
Tachyglossus acu/eatus 116

21

TIGER CAT

Dasyurus maculatus

A, B Cross-sections of various hairs, Maximum diameter of

primary guard hairs up to 11Oj!t-,

C, D Whole mounts of primary guard hairs in shield region.

Whole mount of primary guard hair in proximal half.
E
P-H Scale pattern at various points. F, shield region; G,
mid-shaft region; H, near base,
",
MOST DIAGNOSTIC FEATURES: Aeriform lattice medulla and
medium pigment distribution.

22

&<>1,
?~~

___2 __3~_4~~5__.6__7~_e~~9--"?MM

23