ARTICLE IN PRESS

Journal of Stored Products Research 43 (2007) 2–32

www.elsevier.com/locate/jspr

Review

Biology, behaviour and environmentally sustainable control of

Tineola bisselliella (Hummel) (Lepidoptera: Tineidae)
P.D. Coxa,, D.B. Pinnigerb
a
Central Science Laboratory, Sand Hutton, York YO41 1LZ, UK
b
83 Westwood Green, Cookham, SL6 9DE, UK
Accepted 22 August 2005

Abstract

This review summarises the information available on the biology, behaviour and economic significance of
the common or webbing clothes moth, Tineola bisselliella (Hummel), currently the most important and
widespread clothes moth pest throughout the world. These moths can cause the loss of irreplacable material
of aesthetic, historic and scientific importance, as well as damaging every-day items such as clothes,
furnishings and other materials prepared from animal fur, wool, feathers and hides. Methods for the
detection and control of this pest are outlined, with particular emphasis on control strategies that are
environmentally sustainable and avoid the use of conventional pesticides and fumigants. Improvements in
storage coupled with targeted use of appropriate control measures will help to reduce pest populations.
However, the continuing problems with this pest highlight the need for improved methods of detection,
prevention and management.
Crown Copyright r 2005 Published by Elsevier Ltd. All rights reserved.

Keywords: Webbing clothes moth; Tineola bisselliella; Behaviour; Biology; Control

Corresponding author. Tel.: +44 1904 462000.

E-mail address: [email protected] (P.D. Cox).

0022-474X/$ - see front matter Crown Copyright r 2005 Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.jspr.2005.08.004
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Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
2. Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
2.1. Nomenclature and systematics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
2.2. Morphology and identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
2.2.1. Eggs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
2.2.2. Larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
2.2.3. Pupae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
2.2.4. Adults. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
2.3. Distribution and origin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
2.4. Economic importance and damage. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
2.5. Spread of infestation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
2.6. Life history . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
2.7. Effects of temperature and humidity on development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2.8. Resistance to cold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.9. Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
3. Behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3.1. Larval spinning and locomotion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3.2. Larvae and semiochemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.3. Adult flight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.4. Adults and semiochemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3.5. Adults and sound production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
3.6. Oviposition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
4. Conventional control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
4.1. Insecticides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
4.2. Fumigants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
5. Environmentally sustainable management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
5.1. Detection and monitoring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
5.2. Exclusion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
5.3. Mechanical action . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
5.4. Extreme temperatures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
5.5. Natural enemies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
5.6. Semiochemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
5.7. Modified atmospheric gases. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
5.8. Irradiation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
6. Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

1. Introduction

Clothes moths have been a serious problem for thousands of years, causing damage to clothes,
furnishings and other items made from animal materials such as hides, fur and wool. Today the
common or webbing clothes moth, Tineola bisselliella (Hummel) is the most important and
widespread clothes moth throughout the world. This review has been prompted by the recent
increase in problems associated with this pest in some countries, and the need for improved
methods of detection and more environmentally sustainable control strategies. The review
summarises the published information on the biology and behaviour of T. bisselliella. Methods of
detection and pest control are outlined, with particular attention being given to alternatives to the
use of conventional pesticides.
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2. Biology

2.1. Nomenclature and systematics

The common or webbing clothes moth was first described by Hummel (1823) as Tinea
bisselliella. It was described later in more detail by Zeller (1852), and placed in a new genus
Tineola by Herrich-Schäffer (1853). In earlier literature the name was often misspelt as Tinea
biselliella (Zeller). It has also been referred to under at least six synonyms, all now rejected
(Robinson and Nielsen, 1993).
Common names for this species include the webbing clothes moth (USA and New Zealand), the
common clothes moth (UK and Australia), la teigne des vêtements (France), Kleidermotte
(Germany), polilla de la ropa (Spain), and trac- a dos tecidos (Portugal).
Tineola bisselliella (Hummel) belongs to the family Tineidae in the superfamily Tineoidea. The
systematic position of the Tineidae, a family of world-wide distribution with about 3500 species,
has been reviewed by Robinson and Nielsen (1993). Recently, species-specific differences found in
the quantitative data on the fatty acid content of extracts of whole moths have provided further
evidence of the close relationship between Tineola and Tinea spp. (Cook et al., 1997). The
Tineidae are mostly small, drab moths whose larvae feed on a variety of substrates; however,
unlike most Lepidoptera, very few species attack growing crops. The genus Tineola contains two
species: T. bisselliella found widespread around the world, and T. anaphacola restricted to tropical
West Africa but of doubtful species separation (Gozmány and Vári, 1973; Robinson and Nielsen,
1993).

2.2. Morphology and identification

2.2.1. Eggs
On average, the eggs of T. bisselliella are 0.5 mm long and 0.3 mm at their widest point. Newly
laid eggs are pearly white and subcylindrical in shape with one end more acutely rounded than the
other. To the unaided eye the egg surface appears smooth but under high magnification it is finely
sculptured with a reticulate pattern of irregular polygons and a micropyle at one end.
Photomicrographs showing the development of the larva within the egg are included in a paper by
Titschack (1941). Chauvin (1977) provided a comparison of the eggs of six tineids, including
T. bisselliella and the case-bearing clothes moth, Tinea pellionella (L.), together with line drawings
and photomicrographs of the egg shell structure and a discussion of their significance for water
conservation.
Arbogast et al. (1980) gave a key to help distinguish between the eggs of T. bisselliella and those
of other stored-product moths, based on external morphology as revealed by the scanning
electron microscope (SEM).

2.2.2. Larvae
On emerging from the egg, the larva is white, translucent and about 1 mm long with a head
capsule around 0.2 mm wide. The mouth parts are modified for biting food and for spinning silk.
Reumuth (1946) described the silk-forming organs as long tubular glands alongside the centre of
the intestine, and the spinneret as a pointed appendage in the middle of the lower jaw. The larva
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seeks out a suitable food substrate and spins a fine silken web or tube around itself which is
extended as the larva feeds and moves. Unlike T. pellionella, the larva does not produce a smooth,
portable case. As the larva grows, the integument and head capsule are shed periodically until the
larva is fully grown and ready to pupate. There are usually five or six larval instars, although
Griswold (1944) recorded one particularly long lived individual with 21, and Titschack (1926)
with 41 instars.
The size of the fully grown larva depends on diet and environmental conditions such as
temperature but it is usually in the range 7.0–9.0 mm long and 1.5–1.9 mm broad (Hinton, 1956)
with a head capsule width of about 0.6 mm (Griswold, 1944). The mature larva has a yellowish
brown head with white body, and can weigh up to 3.0 mg (Mallis, 1969). It has three pairs of
jointed thoracic legs, three pairs of prolegs, and an anal clasper on the last segment. There are no
ocelli present on the head. A detailed SEM study of the external morphology and ultrastructure of
chemoreceptor organs has been conducted by Sinitsina and Zaitseva (1988).
A description and illustrated key to larval tineids of economic importance was given by Hinton
(1956) who used morphological characters such as the position of setal hairs on body segments to
identify the different species. A simpler key to European tineids was provided by Carter (1984).
Mound (1989) and Gorham (1991) include T. bisselliella in keys to larvae of moths associated with
stored food in Britain and the USA, respectively. Photomicrographs illustrating the morphology
of the larval skin were included in the publication by Chauvin (1977).

2.2.3. Pupae
When fully grown, the larva stops feeding and usually spins a tough, spindle-shaped, silken
cocoon in which to pupate. The cocoon is smooth on the inside but may be covered with faecal
pellets and food debris on the outside. The size of the pupa is usually in the range 4.0–7.0 mm in
length and 0.8–1.2 mm across the widest point (Griswold, 1944). Weight varies from 3.0–10.0 mg
(Titschack, 1926). As the pupa develops it changes from a creamy white to a yellowish brown on
the head and thorax with black eyes while the abdomen turns a pale tan colour. The tips of the
appendages are free from the body. The posterior abdominal segments are mobile and will wave
about when touched. Each of the abdominal segments 4–9 bears small spines on the dorsal side,
while segment 10 has a small conical projection on each side between which lies the genital
opening. When the adult is ready to emerge, the pupa wriggles from its cocoon prior to ecdysis,
the spines preventing it from slipping back. After the adult emerges, the pupal exuvium is left
projecting from the empty cocoon. Photographs of the pupa and cocoon are included in
publications by Griswold (1944), Pereira (1960) and Chauvin (1977).
Mosher (1916) provided a pupal key to families of the Lepidoptera including the Tineidae,
based on North American species. Mineeva (1974) supplied drawings to help distinguish between
the sexes. Male pupae have distinct boundaries between the last three segments and the genital
opening has a raised tubercle on either side whereas the female terminal segments are less distinct
and the genital opening is smoother.

2.2.4. Adults
The adults vary in length from about 5 to 8 mm with a wingspan of between 9 and 16 mm. The
wings are held tent-like at an angle over the abdomen when the moth is at rest. Unlike
T. pellionella, the wings are without spots or distinct patterns and are bordered by a fringe of long
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hairs. The forewing is pale ochreous with a golden sheen while the hindwing is a shiny pale grey.
SEM photomicrographs of the long slender wing scales have been published in Robinson and
Nielsen (1993). The body has a dense covering of ochreous brown scales, those on the head being
longer and a golden to reddish brown. The compound eyes are black, and the mouth parts are
reduced and functionless. The antennae are threadlike and about three-quarters the length of the
hindwing. The morphology of the mouth parts has been the subject of SEM studies by Faucheux
and Chauvin (1979). The distribution and morphology of the antennal sensilla have been
examined by Faucheux (1985) and Sinitsina and Zaitseva (1988).
Identification of tineid adults to species usually requires dissolving of soft tissues and dissection
of the abdomen so that the structure of the genitalia can be examined (Robinson, 1976). Pierce
and Metcalfe (1935) provided drawings of the genitalia of the British tineids. Photomicrographs
of the dissected genitalia and a key to adults of Australian tineids, including T. bisselliella, were
given by Robinson and Nielsen (1993). Corbet and Tams (1943) and Gorham (1991) prepared
keys to the Lepidoptera infesting stored food products, including T. bisselliella, based on
characteristics of wings and genitalia. Coloured drawings of the adults of all British tineids and a
key based on characteristics of the wings and genitalia were published by Pelham-Clinton (1985).
Other useful works that deal with the identification of tineids include Zagulyaev (1960), Petersen
(1969) and Hannemann (1977).
Recently, differences in the levels of the three main fatty acids found in extracts of whole moths
have been used to distinguish between T. bisselliella and species from other tineid subfamilies,
namely the Tineinae, the Nemapogoninae and the Scardiinae (Cook et al., 1997).

2.3. Distribution and origin

Tineola bisselliella is widespread throughout the world, although Howe and Freeman (1955)
considered it incapable of living in the tropics. The cosmopolitan distribution of this species is
demonstrated by the published records from many different countries, including Australia
(Robinson and Nielsen, 1993), Canada (Monteith, 1973; Sinha and Watters, 1985), Egypt
(El-Sawaf et al., 1967), Europe (Hinton, 1956; Petersen, 1969; Lindroth et al., 1973), New Zealand
(Cox, 1986), Russia (Zagulyaev, 1960), South East Asia (Kawamoto, 1975), Thailand
(Aranyanak, 1993), the USA (Griswold, 1944; Ebeling, 1975) and Zimbabwe (Gozmány and
Vári, 1973). Pelham-Clinton (1985) has prepared distribution maps for this species in the British
Isles, based largely on literature records, and notes that it is less common than it was formerly.
The likely spread of this pest by early sea travel is illustrated by the diary of a naturalist on
board HMS Challenger, during its circumnavigation of the world, who wrote ‘Clothes moths were
a terrible pest and destroyed several garments for me in my cabin.’ (Moseley, 1879). More
recently, records of insects intercepted in Britain on ships’ cargoes between 1957 and 1977 have
been summarised by Aitken (1984); T. bisselliella was found on imports from Australia, Canada,
Russia, South Africa and the USA, infesting products such as dried blood, feathers, bone meal,
mohair, sheepskins and camel hides. It was even recorded on whalemeat in a whaling factory ship.
As well as occurring in domestic premises, museums, grain stores and other buildings,
T. bisselliella has often been recorded in birds’ nests (Linsley, 1946; Woodroffe, 1953; Hicks, 1959,
1962). Nests and other keratin-rich sources of food, such as fur and debris in animal lairs, insect
fragments in bees’ and wasps’ nests and under tree bark (Linsley, 1946), could well have been the
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original habitat of clothes moths before they invaded man-made environments. Tineola spp. may
have originated in southern Africa before being spread throughout the world by human agency
(Meyrick, 1927). However, in a more recent reassessment, Robinson and Nielsen (1993)
concluded that the origin and natural habitat of this species is still obscure.

2.4. Economic importance and damage

Clothes moths have been causing problems for thousands of years, feeding on and damaging
clothes, furnishings, books, art works and other items prepared from animal materials such as fur,
wool, feathers and hides. The main reason for their pest status lies in their unusual ability to digest
keratin. This is the principal protein in these materials which are generally unexploited as far as
other insects are concerned. Their larval midguts have highly reducing conditions, which can
cleave the tightly cross-linked disulphide bonds in keratin to allow a range of proteolytic enzymes
to hydrolyse this otherwise insoluble protein (Hinton, 1956; Waterhouse, 1958; Ward, 1972;
Gleeson et al., 1976).
Tineola bisselliella larvae can also feed on other dry protein-rich materials such as fishmeal,
soya bean meal, dried meat extracts, drugs containing albumin, insect remains, and occasionally
even cereal products and other plant seeds (Cox and Simms, 1978; Aitken, 1984; Carter, 1984;
Stejskal and Horák, 1999). They have occasionally been found infesting mummified human
corpses (Bourel et al., 2000).
The seriousness with which clothes moths have been viewed can be judged by the frequent
references to them in early literature, particularly as symbols of decay. Some of the earliest
references are to be found in both the Old and New Testaments of the Bible, such as in Isaiah
51v.8 ‘For the moth will eat them up like a garment; the worm will devour them like wool.’
(ca. BC 740–540) and in Matthew 6v.19 ‘Do not store up for yourself treasures on earth, where
moth and rust destroyy’ (ca. AD 50–100). References to clothes moths in early Greek and
Roman literature are discussed by Beavis (1988), including examples from Aristophanes (ca. BC
444–385) who describes the moths attacking the feather plumes of helmets, and Pliny (ca. AD 77)
who believed that fabrics used at funerals were immune to attack from clothes moths. Some other
early references are mentioned by Robinson (1979), including those in one of the first printed
books by Joannes de Cuba published in 1491, and the first printed account of the moth in the
English language which appeared in 1521. The French naturalist de Réaumer published a book on
entomology in 1737, on the title page of which was a woodcut depicting early methods of
controlling clothes moths by combing, brushing and fumigation of infested materials.
Early references to this pest in North America have been reviewed by Griswold (1944) who
quotes the travel writer Kalm as saying that clothes moths were troublesome there before the
1750s. They were recognised as museum pests in the USA in the 19th century, damaging insect
specimens for example (Webster, 1892). In New Zealand, Meyrick (1888) noted larvae of
T. bisselliella feeding on chair upholstery.
It is difficult to find reliable estimates of the actual financial damage attributable to clothes
moths. In Britain, official estimates of the cost of damage by textile pests just after the second
world war were put at £1.5 million in 1948, with 80% of all households surveyed stating that they
took some precautions against clothes moths (Hickin, 1974). Parkin (1960) estimated an overall
cost of £3–4 million when pest control measures were taken into account. A decade later, this had
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risen to well in excess of £5 million, and T. bisselliella was acknowledged to be the most common
source of insect damage to textiles (Cornwell, 1971; Tyrer, 1972). By the middle of the 1970s
clothes moths appeared to be less common in Britain and Denmark than previously, possibly in
part due to the increasing use of synthetic fibres (Halstead, 1975; Hallas et al., 1977; Sterling,
1986). Griswold (1944) and Pence (1958) have pointed out, however, that synthetic fibres are not
entirely immune from attack, particularly if they are stained with contaminants such as sweat and
food particles. A photograph of a piece of nylon stocking damaged by larvae of T. bisselliella is
given in the paper by Griswold (1944). More recently, fabrics composed of natural materials
mixed with synthetic fibres have become popular (Bry, 1991). Since T. bisselliella larvae are quite
capable of selecting out and feeding on the wool fibres in these mixtures (Bry et al., 1982), this pest
may well start to increase again. In the USA, where it is found in every state, insect damage to
wool-containing textiles was estimated to cost US$200 million annually in the 1980s (Anonymous,
1981; Reagan, 1982). By the 1990s, Metcalf and Metcalf (1994) suggested that each year
depreciation caused by keratophagous insects including T. bisselliella came to around US$1
billion in the USA alone.
There have been no comprehensive surveys of the current pest status of clothes moths.
Nevertheless they are still very much a problem in areas such as museums and historic houses
where material of aesthetic, historic and scientific importance can be damaged or lost forever
(Florian, 1987; Pinniger, 1991a; Child and Pinniger, 1994; Xavier-Rowe and Pinniger, 2001). In
response to a questionnaire sent to around 100 museums in the British Isles, nearly half the
museums had experienced problems with moths (Linnie, 1987). Materials frequently attacked by
clothes moths in museums include hair, fur, feathers, mammal and bird skins, wool, soiled silk,
upholstered furniture, piano felt, lint and natural bristles, as well as garments and textiles,
particularly those soiled with food, sweat or urine (Zycherman and Schrock, 1988; Pinniger,
2001). Woollen cloth book bindings, paper, imperfectly cleaned animal skeletons, mummified
cadavers and insect specimens have also been damaged by T. bisselliella (Hickin, 1985). A well-
established and widespread infestation was discovered during a survey of a large, five storey
theatrical costume store in the UK which had caused extensive damage to some items (Cox and
Pinniger, 1994). In annual surveys conducted by one of the authors of this review, a number of
museums and historic houses in the UK have experienced serious infestations over the last 5 years;
several major national museums and a leading art gallery have reported two generations a year
recently, with warmer winters and wetter summers possible contributory factors.
In other parts of the world, clothes moths also continue to cause serious concern, for example in
major wool-producing countries such as Australia and New Zealand (Mackerras, 1970; Waller,
1984). In Canada, T. bisselliella is also occasionally found in grain elevators, although the moths
could have been associated with the nests of birds or rodents in the grain elevators (Sinha and
Watters, 1985).
It is the larvae of clothes moths that cause all the damage, the adults being unable to feed.
Typical damage takes the form of holes, irregular in shape and size, bitten through the fabric or of
surface browsing by biting through the bases of raised fibres (Parkin, 1960). For example, larvae
will chew through the warp threads of a wool carpet so that the pile falls out (Moncrieff, 1950).
Only a small amount of feeding will ruin materials such as clothing.
Larvae cause further damage by producing large numbers of webbing tubes and sheets
containing considerable quantities of faecal pellets which spoil the appearance of the product.
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Damage is often concentrated on soiled areas of materials, and in dark folds and creases
(Fraenkel and Blewett, 1946; Pinniger, 1991a). Materials are more likely to be attacked if they
have been left undisturbed for a long time in unlit conditions, such as in store cupboards, under
heavy furniture that is rarely moved and standing on carpets, and in fluff accumulating in cracks
between floorboards (Griswold, 1944). Large larvae will chew holes even in materials which they
cannot digest, although cotton is rarely attacked. Young larvae, on the other hand, may not be
able to attack coarse animal hairs because their mandibles are too small and can only span about
90 mm (Busvine, 1980).
Photographs of damage caused by T. bisselliella have been published by Titschack (1941),
Griswold (1944), Pereira (1960), Mourier and Winding (1975), and Robinson and Nielsen (1993).
Photomicrographs of the marks left by larval mandibles on wool fibres, on plastic bags, and on
lettuce seeds can be found in a publications by Reumuth and Loske (1960), McPhee (1971), and
Stejskal and Horák (1999), respectively. Microscopic examination of the damage to wool fibres
caused by digestion in the larval gut has been undertaken by numerous workers, including Day
(1951), and Hammers et al. (1987). Gerard and Ruf (2002) found that wool samples with high
fibre curvature had lower mass losses from larval feeding damage than wool samples with low
fibre curvature. Behavioural responses of T. bisselliella larvae to staple structure, including wool
characteristics such as fibre curvature and density, were considered to be responsible for the
significant wool mass loss differences caused by insect feeding between fine and coarse wools.
When staple structure was destroyed through fine chopping, the degree of fibre curvature and
bulk prior to chopping influenced the extent of larval moulting.

2.5. Spread of infestation

Clean material can become infested by T. bisselliella adults flying in from outside sources such
as birds’ nests in roof spaces or moving from other infested materials nearby. Even dead mice may
provide a source of infestation in buildings (Salmond, 1956). Similarly, textiles can become
infested by larvae crawling in from surrounding areas; newly hatched larvae can pass through
crevices that are only 0.01 mm in width (Colman, 1940). Clothes moths can also be introduced
when new materials that have already been infested with eggs laid elsewhere are brought into
store. For example, packaging and display materials can be a source of infestation in museums
(Pinniger, 2001). The origins and types of harbourages for insects in museums have been reviewed
by Pinniger and Winsor (1998). Imports may become cross-infested from residues left in ships’
holds and containers from previous cargoes (Aitken, 1984).

2.6. Life history

Unlike insects feeding on growing plants, the life history of the clothes moth is not constrained
by the seasonal availability of their food supply. To some extent they are also buffered from
seasonal variations in temperature by spending most of their life cycle in protected habitats such
as buildings. Nevertheless, temperature, relative humidity, type and availability of food remain
the chief factors affecting the duration of the clothes moth’s life cycle. Indoors, development is
continuous and T. bisselliella passes through numerous generations each year. However, in birds’
nests and other exposed habitats development may be prolonged by cold and lack of food, with
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larvae going through many more than the usual five or six instars (Hinton, 1956). Zagulyaev
(1954) states that this species is univoltine in some parts of Russia.
Some authors have suggested that T. bisselliella larvae may also undergo a diapause under
certain adverse conditions, although this has never been confirmed in the laboratory (Hinton,
1956; Pelham-Clinton, 1985). Diapause is a state of delayed development or dormancy which is
usually closely linked to the insect’s natural environment, enabling it to survive periods of extreme
conditions such as cold or food shortage. In other storage moths, such as the brown house moth,
Hofmannophila pseudospretella (Stainton), the warehouse moth, Ephestia elutella (Hübner), and
the Mediterranean flour moth, E. kuehniella Zeller, diapause occurs in the last larval instar before
pupation and often helps to synchronise subsequent adult emergence (Woodroffe, 1951; Cole and
Cox, 1981; Bell, 1989). Daylength has been shown to play an important role in diapause induction
in storage moths (Cox, 1975; Bell, 1976; Cox et al., 1981) but its effects on T. bisselliella have yet
to be studied in detail.

2.7. Effects of temperature and humidity on development

There are no comprehensive developmental data published for T. bisselliella at different

combinations of temperature and relative humidity compared to other stored-product moths such
as the tropical warehouse moth, E. cautella (Walker), and E. kuehniella (Burges and Haskins,
1965; Jacob and Cox, 1977). The little information that has been published is scattered
throughout the literature, and the more reliable data are summarised in Table 1. On poorer diets,
such as feathers and human hair, development can take well over a year to be completed
(Griswold, 1944).
Most larvae are able to complete development down to 30% r.h. (Griswold and Crowell, 1936).
Relative humidity had little effect on the length of the egg stage in the range 20–100% r.h. at
13–32 1C (Griswold, 1944). Hinton (1956) has suggested that T. bisselliella can survive conditions

Table 1
The effect of temperature and relative humidity on the development of Tineola bisselliella

Temp (1C) r.h. (%) Food Stage Days Reference

23.5 50 Impregnated flannel E–A 63 Pereira (1960)

31.0 66 Impregnated flannel E–A 55 Pereira (1960)
24.0 75 Fishmeal E–A(F) 49 Griswold and Crowell (1936)
24.0 75 Fishmeal E–A(M) 42 Griswold and Crowell (1936)
25.0 70 Soya bean meal E–A 66 Cox and Simms (1978)
20.0 55 Impregnated flannel O–Eh 10 Pereira (1960)
25.0 70 Impregnated flannel O–Eh 7 Pereira (1960)
33.0 64 Impregnated flannel O–Eh 7 Pereira (1960)
13.0 20–100 Fishmeal O–Eh 37 Griswold (1944)
18.5 20–100 Fishmeal O–Eh 14 Griswold (1944)
24.0 20–100 P 12 Griswold (1944)
13.0 20–100 P 51 Griswold (1944)

A ¼ adult, E ¼ egg, Eh ¼ egg hatch, F ¼ female, M ¼ male, O ¼ oviposition, P ¼ pupa.

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of low humidity because of its ability to utilise metabolic water produced as a by-product of the
digestion of keratin. Generally, the life cycle is faster at higher relative humidities (Titschack,
1925; Griswold, 1944; Fraenkel and Blewett, 1946; Whitfield and Cole, 1958; Chauvin, 1977). The
life history of T. bisselliella in an Australian bulk wool store has been described by Key and
Common (1959).
Development and reproduction can occur at 33 1C, and eggs hatch at 35 1C (Rawle, 1951).
However, adult males become sterile within 2 days if kept at 35 1C, and all life stages are killed
within 4 h at 41 1C (Rawle, 1951). When larvae were reared on a diet of fishmeal, mean longevity
of the resulting mated adult males ranged from 38 days at 18.5 1C to 29 days at 24 1C, and female
longevity from 22 to 13 days at the same temperatures, respectively (Griswold, 1944). However,
according to El-Sawaf et al. (1967) who reared larvae on dried meat or wool fibres, at 25 1C and
100% r.h. mated females lived about 6 days but only 4 days at 31 1C 60% r.h.; unmated females
lived on average 12 days under these conditions. At 80% r.h. about 100 eggs were laid per mated
female at 30 1C compared to only 65 eggs at 25 1C; unmated females produced only around 33
eggs each (El-Sawaf et al., 1967).

2.8. Resistance to cold

Solomon and Adamson (1955) classified this species as cold hardy when they found it could
survive a British winter in unheated conditions during which temperatures dropped to 0 1C or
lower on 75 days. According to Howard and Marlatt (1896), larval feeding ceases at 9 1C and
larval movement stops at 4–6 1C. Back and Cotton (1926) reported that larvae of T. bisselliella
survive temperatures of 6.7 to 3.9 1C for 67 days and 15 to 12.2 1C for 21 days. Mature
larvae survived 1 year at 4 1C (Back and Cotton, 1927). More recently, Brokerhof et al. (1993)
found that at 20 1C the longest times with survival were 16 h for eggs but less than 1 h for larvae,
pupae and adults. The 0–2 day old egg was the stage most tolerant to low temperatures, some
survival occurring after 2 days exposure to 15 1C, 3 days at 10 1C, 7 days at 5 1C and 21 days
at 0 1C (Brokerhof et al., 1992).

2.9. Nutrition

Early studies on the physiology of nutrition, including digestion, excretion and water balance,
have been summarised by Wigglesworth (1972). More recent work on aspects of the nutritional
ecology, including food utilisation and relative growth rates, of keratinolytic insects such as
T. bisselliella have been reviewed by Baker and Bry (1987).
Feeding is continuous throughout the larval period, except during moulting between instars.
Illingworth (1917) claimed that the larvae can become cannibalistic when food is scarce, although
Griswold (1944) and Whitfield and Cole (1958) found no evidence for this in their own studies.
Studies on wool digestion in T. bisselliella have shown that before keratin can be attacked
successfully by proteinase enzymes, the disulphide bonds have to be broken apart by reduction of
cysteine to cystine and water under the anaerobic conditions found in the larval midgut (Powning
and Irzykiewicz, 1962). After this, digestion is completed by a mixture of amino- and
carboxypeptidases. Ward (1972) noted that the many different proteolytic enzymes present were
unlikely to be the products of gut microflora, particularly since Crewther and McQuade (1955)
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found only low numbers of gut bacteria in T. bisselliella. The central position of cystine in the
excretory mechanism seems to be confirmed by the high levels (10% of dry weight) of this
substance in excreta (Powning, 1953). However, Wigglesworth (1972) suggested that these high
levels may be the result of cystine-rich fragments of undigested wool present in the faecal pellets
because wool contains high levels of sulphur amino acids.
More recent studies on the digestion of wool in T. bisselliella have been reviewed by Robinson
et al. (1993). The midgut proteinase enzymes of T. bisselliella have been characterised by
Christeller et al. (1994). They demonstrated that the major endopeptidase activities were the result
of serine proteinases which exhibited hydrolytic activity against N-benzoyl-DL-arginine-p-
nitroanilide and N-succinyl-L-alanyl-L-alanyl-L-prolyl-L-leucine-p-nitroanilide.
Most of the studies on nutrition in T. bisselliella have centred on its unusual ability to digest
keratin. However, this species is by no means restricted to such diets, and Fraenkel and Blewett
(1946) have examined its more general food requirements. They found that this was one of the few
insects tested in which carbohydrates did not appear to be necessary for larval development.
Vitamins of the B complex were essential for successful growth; little or no growth took place
when pantothenic acid or nicotinic acid were absent, and growth was retarded in the absence of
riboflavin, choline chloride, pyridoxin or thiamin. Inositol and p-amino benzoic acid did not
appear to be necessary for development, and the value of biotin was considered doubtful. This
agrees with observations by many other workers that T. bisselliella cannot survive on thoroughly
clean wool alone and, indeed, develops faster and more successfully on other foodstuffs
containing nutrients such as the B vitamins which are missing from keratin fibres (Titschack,
1922; Crowell and McKay, 1937; Griswold, 1944; Pereira, 1960).
Griswold (1944) tested 27 different foodstuffs and found that fishmeal was the most satisfactory
for rearing T. bisselliella in the laboratory. At the UK Central Science Laboratory (CSL), clothes
moths have been reared successfully for many years on a diet of fishmeal and dried yeast powder
(16:1 by weight) to which is added a piece of felt material containing at least 35% wool fibres. In
the Savannah USDA laboratory, wool cloth sprinkled with a little brewer’s yeast is used for this
purpose, as specified in the US CSMA textile resistance clothes moth test method (Anonymous,
1971). Other laboratory methods have included cholesterol and casein in the diet (Anonymous,
1978). Whitfield and Cole (1958) found that T. bisselliella developed fastest on a diet of flannel,
yeast and cholesterol.

3. Behaviour

3.1. Larval spinning and locomotion

One of the main characteristics of larval behaviour in the family Tineidae is that of silk spinning
in the form of cases, tunnels and random webbing of the substrate. This activity may help to
conceal as well as physically protect larvae from attack by parasites and predators (Scoble, 1992;
Robinson and Nielsen, 1993) and may even slow down desiccation in dry conditions. Thus, at 0%
r.h. water loss in T. pellionella larvae without cases was twice that of larvae enclosed in cases
(Chauvin et al., 1979). Unlike many other clothes moths, T. bisselliella does not build a case but
spins a rather crude tunnel whilst moving through its food. It also meshes together food particles,
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faecal pellets, cast skins and other materials wherever it moves, behaviour which has given rise to
one of its common names of ‘webbing clothes moth’. It has been estimated that each larva
produces about 0.1 mg of silk per week, that is 1.7% of its mean body weight (Busvine, 1980).
These activities have been likened to modifications of cocoon-spinning prior to pupation
(Robinson and Nielsen, 1993). Larvae usually remain within their feeding tunnels, although if the
food becomes unsuitable they may leave them in search of fresh supplies (Hinton, 1956). Larvae
also produce tough cocoons in which to moult and pupate (Griswold, 1944).
Although larvae have no ocelli, they are able to distinguish between light and dark. The larvae are
negatively phototactic and when placed on a piece of material they will crawl to the darkened
underside as quickly as possible (Griswold, 1944). They seem to prefer to live between layers of
material or within powdered foodstuffs so that they are surrounded by the substrate and hidden
from view. General observations also suggest that larvae are sensitive to and disturbed by vibration.
Little information has been published on the movement of larvae to new sources of food,
although it has generally been assumed that larvae will remain with a suitable food source until it
is exhausted. In some unpublished work at CSL, 0–1 day old larvae crawled towards raw sheep’s
wool placed 15 cm from the larvae but not when the wool was 23 cm or more away; older larvae
moved very little from their original points of infestation in a small room when plenty of food was
available. Mature larvae prefer to migrate to and pupate at the outer surface of the feeding
material or in creases and folds, a characteristic confirmed during field studies in an Australian
wool store by Key and Common (1959). Lines of pupal cases are often found positioned at the
edges of heavy infestation sources, thus facilitating the subsequent emergence and dispersal of the
moths.

3.2. Larvae and semiochemicals

Semiochemicals have been defined as chemicals that mediate interactions between organisms
(Law and Regnier, 1971); their mode of action and potential for use to control insect pests of
stored products have been reviewed recently by Cox (2004). These chemicals can be produced
either by the insects themselves or by other organisms, including the plants upon which they feed,
and are important because of their central role in the behaviour of insects.
There is little information published on the effects of semiochemicals on larval behaviour.
Observations by Mallis et al. (1959, 1962), Bry (1991) and others have confirmed that fabrics
contaminated with sweat, urine and food stains are more commonly damaged by larvae than are
clean fabrics. Although foodstuffs have been ranked for feeding damage or their suitability for
larval development, few choice tests have been conducted. However, unpublished work at CSL
showed that when feeding larvae were given a choice, they preferred flannel, feather, fur and flock
to horsehair or paper. Pereira (1960) found that flannel impregnated with yeast and cholesterol
was the most preferred food for larvae, followed by rabbit skin, knitting wool and woollen cloth
in decreasing order of preference.

3.3. Adult flight

In common with other moths, on emergence from the pupal case the wings of T. bisselliella
are small and crumpled. Fluid is pumped through the veins while the wings are held vertically
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(Roth and Willis, 1952). The wings are then brought down parallel with the abdomen to
dry; after about an hour they reach their full size and the moths are ready for flight (Hickin,
1985).
Adults are negatively phototactic and will usually fly or crawl to a darkened area to hide away
from light as rapidly as possible (Griswold, 1944). It has been claimed that T. bisselliella can fly up
to 90 m to a new source of food (Herrich, 1933). It is usually only the males that take wing, the
females preferring to move with a characteristic scuttling run (Key and Common, 1959). Flight is
more common when it is hot, although the minimum temperature for flight is unknown. Moths
can run at speeds around 10–13 mm/s compared to a flight speed of 50–70 mm/s (Busvine, 1980).
Moths will also sometimes move along by a series of rapid but short hops, using the hind legs and
a few beats of the wings.
Field studies using sticky traps in wool stores indicated there was a peak of flight activity
around dusk, with about half the moths being caught during the 3 h after sunset (Key and
Common, 1959). They also found that a rise from about 18 to 27 1C doubled the catch.
Laboratory studies by Chauvin (1977) showed that at 20 1C and 50% r.h. flight only took place at
times corresponding to the hours of natural darkness between dusk and dawn, even when the
moths were kept in continuous darkness.

3.4. Adults and semiochemicals

Mating behaviour in laboratory cultures has been described by Titschack (1922), Roth and
Willis (1952) and Zagulyaev (1960). Photographs of courtship and mating were reproduced in the
paper by Roth and Willis (1952). Mating can occur within a few hours of emergence from the
pupal case and may last for up to 1 h but more usually for about 15 min. Females emit a
pheromone by slightly raising the end of the abdomen above the wings and extruding the tip of
the ovipositor. When virgin females were kept on a sheet of filter paper, the pheromone was
apparently imparted to the paper and could be extracted with petroleum ether (Roth and Willis,
1952). The crude extracts could be kept for a year at low temperatures and still retain their ability
to attract males. The sexually active male walks about rapidly and erratically, continually
vibrating the wings. Wing vibration continues as the male approaches a female which remains
more or less stationary. During copulation the male grasps the female genitalia with his claspers
and moves round the female until they are in a linear position with heads pointing in opposite
directions.
Yamaoka et al. (1981, 1985) used capillary gas-liquid chromatography and mass spectroscopy
to determine the structures of pheromones produced by female T. bisselliella. So far, two
compounds have been identified; they have been named as Koiganal I and II, and appear to be
produced in the ratio 1:2, respectively. Koiganal I has been identified as (2E)-octadecenal and
Koiganal II as (2E,13Z)-octadecadienal. The synthesis of both compounds has been described by
Takikawa et al. (1997).
Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass
spectrometric (GC-MS) analyses, Takacs et al. (2001c) have identified two compounds from
methanolic extracts from male T. bisselliella, which appear to have aggregation properties. These
are methyl hexadecanoate and methyl (9Z)-hexadecenoate. In bioassay arena tests, a blend of
these two esters was attractive to male and virgin female moths but not mated females.
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In choice arena bioassays, Takacs et al. (2001a) provided evidence that long-range attraction to
suitable larval habitats for mating and egg-laying purposes was mediated by semiochemicals.
Male and female adult T. bisselliella were attracted to dried but untanned animal pelts and
preserved horseshoe crab but not to unprocessed sheep’s wool. Volatiles were identified by
comparative GC-MS and GC-EAD analyses, and a blend of the aldehyde, nonanal, and
geranylacetone appeared to be the most attractive components from 28 candidates (Takacs et al.,
2001b).
The uses of semiochemicals for detection and control are considered in Sections 5.1 and 5.6,
respectively.

3.5. Adults and sound production

Sound production has been reported to be part of the communication system of several stored-
product moths. Ultrasound production has been demonstrated in the lesser wax moth, Achroia
grisella (F.), the rice moth, Corcyra cephalonica (Stainton), the Indian meal moth, Plodia
interpunctella (Hübner), E. cautella, and E. kuehniella (Spangler et al., 1984; Spangler, 1987;
Trematerra and Pavan, 1995). Low-frequency communication signals have been reported in the
greater wax moth, Galleria mellonella (L.) (Spangler, 1985).
Low-frequency signals have also been implicated in the communication systems of adult T.
bisselliella (Takacs et al., 2003). Males produced sounds of 27 dB with a base frequency of
40–50 Hz and a harmonic frequency of 80–100 Hz. Sound intensity and frequency increased to
55 dB and 65–75 Hz, respectively, when calling males were less than 2 cm away from conspecifics
of either sex. In Y-tube bioassays, virgin male and female moths preferred played-back sonic
signals from males to a silent control, whereas mated females showed no preference for either
stimulus. Use of pheromonal as well as sonic signals by T. bisselliella may be adaptive in ensuring
successful communication in enclosed microhabitats. Should prolonged exposure to male-
produced aggregation pheromone or female-produced sex pheromone (Takacs et al., 2001c) in a
confined environment cause adaptation or habituation of the olfactory system, males and females
should still be able to respond to sonic signals.

3.6. Oviposition

Eggs can be laid singly or in groups by both virgin and mated females, and oviposition usually
commences within 24 h of mating (Griswold, 1944). Kan and Waku (1985) observed that at 25 1C
and 45–50% r.h. the first egg is laid 3–7 h after mating. The long soft ovipositor is extended
telescopically and inserted into a space to test its suitability before laying an egg, usually among
the fibres of fabric or particles of other food materials. In the absence of larval food, eggs can also
be laid on non-food materials such as in the crevices between overlapping sheets of paper. This
behaviour has been used as a laboratory technique for collecting eggs of this species (Pence, 1958).
The eggs are not glued very strongly to the substrate and can easily be dislodged (Hinton, 1956).
Analysis of oviposition preferences in T. bisselliella has been undertaken by Kan and Waku
(1985) who also presented an SEM of the ovipositor revealing the presence of numerous sensory
hairs. The ultrastructure of the receptors on the ovipositor of this species has been examined by
Faucheux (1988). Tactile stimuli seemed to be more important than chemical stimuli for
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oviposition control. Using wire gauze of various mesh sizes, Kan and Waku (1985) found that the
females preferred to oviposit on 40 mesh gauze with a distance between the wires of 0.45 mm.
They also found that eggs were laid preferentially on fleecy substrates irrespective of whether they
were wool or cotton.
In laboratory choice tests, Traynier et al. (1994) observed that odours from dried yeast localised
oviposition on fabric. This effect was augmented by odours from the faeces of conspecific larvae
that had eaten a favourable diet containing dried yeast. Some plant phenolics, flavonoids and
quinones incompletely deterred oviposition; both vegetable oils and fatty acids from wool wax
deterred oviposition equally well when present in large amounts. Studies by other workers on
oviposition preferences have been rather contradictory. Similarly, investigations of the effects of
temperature and relative humidity on oviposition were inconclusive (El-Sawaf et al., 1967).
The number of eggs laid by each female is variable but is usually about 100, depending on
factors such as the weight of the female. Benedict (1917) counted between 30 and 160 eggs per
female, while Titschack (1922) recorded a maximum of 221. The maximum number of eggs laid in
1 day by a female was 59 (Back and Cotton, 1931). At 25–30 1C fewer eggs were laid at 60% r.h.
than at higher humidities (El-Sawaf et al., 1967). Unmated females lay fewer eggs than mated ones
(Pereira, 1960). The length of the oviposition period is very variable and is longer at lower
temperatures. Thus, Griswold (1944) recorded a mean of 17 days (range 8–27) at 18.5 1C
compared to only 11 days (range 5–21) at 24 1C; however, there was little difference in the total
number of eggs laid when adults were kept at these different temperatures. Most females die soon
after ceasing to lay eggs.

4. Conventional control

4.1. Insecticides

The conventional method of preventing and controlling clothes infestations is by the use of
insecticides, the synthetic pyrethroids permethrin and cypermethrin being the main active
ingredients of modern moth-proofers. For example, in many countries woollen carpets are
regularly treated with permethrin during manufacture, usually at the yarn-dyeing stage, which
provides long-term protection against attack (www.canesis.com/Documents/Yarn_Chemical_
Treatments.pdf, last accessed 9 December 2005). Another synthetic pyrethroid, vaporthrin
(empenthrin), has also been shown to be effective against clothes moths (Yoshida et al., 1984;
Chio et al., 1996).
The use of chemicals to control clothes moths and other textile pests has been reviewed by
Mallis (1969), McPhee (1971) and Reagan (1982). Chemical pest control of insects, including
clothes moths, in museums has been summarised by Linnie (1987), Pinniger (1991b, 2001),
Pinniger and Child (1996), Pinniger and Winsor (1998), and Pinniger and Harmon (1999).
A microemulsion formulation of permethrin has shown promise for the control of T. bisselliella in
museums with minimum impact on materials (Pinniger et al., 1994), as has an aerosol formulation
of another pyrethroid, cyphenothrin (Webb et al., 1994). However, pesticide registrations are
continually changing due to health and environmental concerns, and the current approval status
of particular chemicals should always be checked with the appropriate registration authority. For
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example, dichlorvos (DDVP) slow-release strips are particularly effective but their use is banned
in many countries, including the UK, because of the associated health risks (Pinniger and Child,
2003).
Standard methods for testing the effectiveness of moth proofing agents are described in
BS4797-ISO3998 (Anonymous, 1978) and the American textile resistance test (Anonymous,
1971).
The main problems associated with the use of conventional insecticides to control clothes moths
are toxicity to non-target organisms in the environment, including humans, and the development
of resistance in the pests. For example, there is a serious risk of pollution when effluent from the
pesticide treatment of carpets is discharged to the environment, particularly where closed-loop
systems are absent. Although it is relatively easy to kill adult moths with low doses of pyrethroid,
the larvae can be very difficult to control. It is almost impossible to deliver an effective dose
of a residual insecticide to a larva hidden inside folds of fabric and protected by its tunnel of
silk webbing. Health problems associated with the use of some of these chemicals have been
reviewed by Linnie (1990). A general account of the biochemistry and genetics of insect resistance
to pesticides is given by Oppenoorth (1985), and information on this subject specifically for
T. bisselliella is discussed by Robinson (1971) and Kerr (1977).

4.2. Fumigants

Some chemicals used as insect repellents, including naphthalene and para-dichlorobenzene, also
have a fumigant action and are toxic in high concentrations (Reagan, 1982). However, the term
‘fumigant’ is usually only applied to toxic gases. The use of fumigants, such as ethylene oxide,
methyl bromide, phosphine and sulphuryl fluoride, for the control of clothes moths is uncommon
except in museums, and has usually been restricted to severe, hidden infestations (Derrick et al.,
1990; Pinniger and Harmon, 1999; Griffin, 2001).
Fumigant gases will penetrate materials and kill all the susceptible insects in the target objects
but the treatment provides no residual protection. Other disadvantages of using fumigants include
the need for specialised equipment and their high toxicity to humans which restricts their use to
fully trained staff only (Zycherman and Schrock, 1988; Pinniger, 2001). Most fumigants have
deleterious effects on metals and other materials such as wool and horsehair (Pinniger and
Harmon, 1999). In addition, methyl bromide has damaging effects on the ozone layer (Price,
1996). Thus, many of the older fumigants are now being abandoned, phased out or made illegal
for all but certain critical uses because of safety and environmental issues.

5. Environmentally sustainable management

Concerns over the adverse effects of the continued use of conventional pesticides and fumigants
on human health and the environment, together with continuing problems of pest resistance,
have encouraged the search for safer and environmentally sustainable alternatives for pest
management. The following sections review the main options currently available for reducing
the use of conventional pesticides and fumigants to prevent or control infestations of clothes
moths.
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5.1. Detection and monitoring

Detection of clothes moths and their immature stages is difficult because of their small size,
drab colour and cryptic behaviour. Inspection methods and where to look for signs of infestation
have been described recently by Zycherman and Schrock (1988), Pinniger and Winsor (1998), and
Pinniger and Harmon (1999).
Although a thorough visual inspection of materials and buildings, both with the unaided eye
and by microscopic examination, is an important first step, the chances of detecting low-level
infestations before they become more serious can be increased significantly by using insect traps.
In some early work, Wilson (1940) found that clothes moths were attracted into box traps
containing cloth pads treated with fishmeal or an alcoholic extract of fishmeal; sticky fly papers
also caught clothes moths, especially when treated with various animal substances such as fish oils
and meat scraps. The use of food volatiles to attract moths to traps may be worth consideration
for further research.
Pinniger (1991b) reported that sticky traps designed for cockroaches have been used
successfully for the detection of T. bisselliella. Now that the sex pheromones of this species
have been synthesised, it has been possible to develop more efficient flight traps containing
attractant lures. These have been tested in trials in a UK theatrical costume store and in a USA
natural history museum where the traps were used successfully to monitor moth populations (Cox
et al., 1996). The results showed that traps with pheromone lures caught about 20 times the
number of moths caught on similar but unbaited traps. Since then, pheromone-baited sticky traps
have been used regularly to monitor T. bisselliella in many situations, particularly in museums and
historic houses (Pinniger, 1995; Carter, 1998; Pinniger et al., 1998; Xavier-Rowe and Pinniger,
2001). They have also been used successfully to monitor this species in a warehouse containing
woollen materials where they were found to work best in warm (23–27 1C) and stagnant air
conditions with low light levels (Trematerra and Fontana, 1996).
The correct selection, positioning and the number of traps to deploy in any given situation is
vital to the accurate identification of problem areas, the targeting of appropriate control measures
and the monitoring of their effectiveness. These aspects of trapping have been described in more
detail by Florian (1987), Pinniger (2001), Child and Pinniger (1994), Pinniger and Harmon (1999),
Pinniger and Child (2003), and Pinniger et al. (2004). However, it should be remembered that
although some of the pheromone lures currently available commercially are extremely effective,
they only attract male moths, and so visual inspection should be used to supplement traps
(Pinniger and Winsor, 1998).

5.2. Exclusion

Wherever possible, items at risk from attack by clothes moths should be stored in air-tight
containers that physically prevent the entry of moths and their larvae, thereby avoiding the need
for toxic chemicals. However, even tiny gaps in the seals may allow the entry of young larvae.
Traditional methods of keeping clothes moths away from susceptible materials have been
reviewed by Mallis (1969). In addition, ‘Gore-tex’ membrane envelopes, made of expanded
polytetrafluoroethylene (PTFE) laminated to a non-woven polyester fabric, have been shown to
have potential for protecting woollen materials from clothes moths (Bry and Mullen, 1988). Many
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museums in the UK successfully protect textiles with ‘Tyvek’, a material composed of very fine,
high-density polyethylene fibres (Pinniger, pers. comm.).

5.3. Mechanical action

Mechanical agitation in the form of activities such as beating, brushing and combing of
materials has been used for many years as a method of destroying the eggs, larvae and pupae of
clothes moths. However, these methods are not effective against the adult stage and can only be
used on more robust materials. Vacuum cleaning will also disinfest contaminated fabrics,
although eggs may be difficult to dislodge if they become trapped between the fibres. Low pressure
has been used to control museum insect pests; for example, larvae of the varied carpet beetle,
Anthrenus verbasci (L.), were killed by 1 day at a constant pressure of 20 hPa (Bergh, 1998).

5.4. Extreme temperatures

In all living animals, the rates at which most physiological activities and associated functions
proceed are related to body temperature. Insects are often described as cold-blooded and are
generally considered to be ectotherms, that is they are unable to regulate their body temperature
metabolically and have to rely on absorbing heat from their surroundings (Heinrich, 1973). Thus,
manipulating the temperature of the environment surrounding an insect offers a non-chemical
route to pest control. The use of extreme temperatures to control stored-product insects generally
has been reviewed recently by Fields and Muir (1995) and Burks et al. (2000).
The optimum temperature for maximum rate of development and population increase lies in
the range 25–33 1C for the major stored-product insect pests, while developments stops altogether
below 10 1C and above 35 1C. (Howe, 1965; Bell, 1975; Fields, 1992). Storage insects can survive
temperatures outside this range depending on various factors such as the exposure period, the
strain and developmental stage, and any acclimation resulting from prior exposure to a less severe
temperature regime.
Death of many species will occur within hours of exposure to 45 1C, while some previously
acclimated insects will require months of exposure to kill them even at 10 1C (Fields, 1992).
Thus, it would be necessary to cool a commodity, currently at a temperature optimum for
development, by nearly three times as many degrees centigrade as it would to heat it to ensure all
infesting insects are killed. However, this is not the only factor to be considered when choosing
between high and low temperatures for control. The effects of extreme temperatures on the
commodity itself need to be assessed; the thickness, density and moisture content of the material
to be disinfested, and the previous history and acclimation of the particular insect strain involved
can all have a significant effect on the exposure period required to ensure complete kill at a
particular temperature.
Death resulting from exposure to high temperature is due to the chemical components of cells
becoming unstable and damaged irreparably, particularly through the denaturing of intracellular
proteins and the disruption of cell membranes (Denlinger et al., 1991). The mechanisms involved
in heat death in insects have been reviewed recently by Fields (1992) and Burks et al. (2000).
Male moths of T. bisselliella are sterile after 2 days at 35 1C; all stages die after only 4 h at 41 1C
or 11 min at 49 1C (Rawle, 1951; Back, 1923). The thermostupor point, that is the temperature
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when heat stupor causes an individual to become motionless immediately before death, has
been calculated as 50 1C (Vannier, 1994). Thus, steam ironing or washing of materials infested
with clothes moths may be effective if the temperature is sufficiently high. The principles of
heat disinfestation as they apply to treating museum collections have been summarised by
Strang (2001). Some objects can be placed in an oven at 52 1C for 24 h if placed in airtight bags;
more sensitive items can be treated in a special humidity-controlled chamber at the same
temperature using the Thermo Lignums process (Pinniger, 1996; Ackery et al., 2001). One to two
hours at the most at this temperature is sufficient to kill the insects, the longer time stated being to
ensure that infested objects are gradually brought up to the required temperature and back down
again.
Death resulting from exposure to low temperatures is due to irreversible mechanical and
osmotic damage to cell membranes as insect body fluids freeze and ice crystals form leading to the
collapse of cells (Lee, 1991). The mechanisms involved have been reviewed in more detail by
Fields (1992) and Burks et al. (2000). Ice crystals start to form around ice nucleators such as food
particles, proteins and alcohols, and in their absence water can remain liquid at temperatures well
below 0 1C. The temperature at which ice crystals begin to form spontaneously in a supercooled
liquid is referred to as the supercooling point (Salt, 1961; Somme, 1982). Smaller insects and
developmental stages such as eggs tend to have lower supercooling points than larger ones
(Somme, 1982). Supercooling points measured so far for storage insects range from 7 to 21 1C
and these temperatures cause death within minutes, although temperatures above them are often
lethal too (Fields, 1992).
All stages of T. bisselliella are killed after 1 day at 18 1C (Back and Cotton, 1926). More
recently, Brokerhof et al. (1992, 1993) have studied time–temperature–mortality relationships
of eggs of T. bisselliella at temperatures in the range 20 to 0 1C, and for all developmental stages
at 20 1C. Eggs were the most tolerant stage and required 30 h exposure at 20 1C for complete
kill. At this temperature, larvae, pupae and adults were all killed after only 1 h. Eggs were also
exposed to a slow cooling rate of 2 1C h1 to a minimum of 20 1C, and no evidence of
acclimation during cooling was found (Brokerhof and Banks, 1993). Chauvin and Vannier
(1997) found the supercooling points for eggs and second instar larvae to be 23 and 13 1C,
respectively.
The widespread use of low temperatures as a method of pest control in museums, together with
precautions to prevent damage, such as sealing materials in airtight polythene bags to avoid
condensation, has been summarised by Stansfield (1989). Pinniger (1991b) noted that in a trial control
programme, mixed age larvae on museum textile objects were all dead after 7 days at 30 1C. Large
collections of ethnographic objects with heavy infestations of T. bisselliella have been successfully
treated in Sweden at 29 1C for 10 days and in Canada using two freeze-thaw cycles down to 18 1C
(Griffin, 2001; Petersson, 1998). However, 3 days at 30 1C or at least 14 days at 18 1C without
cycled freezing and thawing will usually kill all stages of this species (Pinniger, 2001).

5.5. Natural enemies

Based mainly on casual observations in the field, larvae of the window fly, Scenopinus fenestralis
(L.) are known to be predaceous on T. bisselliella, sucking the body fluids from larvae (Griswold,
1944; Hinton, 1956). Other predators include the spiders, Uloborus spp., and mites of the genera
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Blattisocius, Pediculoides, Pymotes and Typhlodromus (Griswold, 1944; Hinton, 1956; Herting,
1975).
There are numerous field records in the literature of hymenopterous parasitoids attacking T.
bisselliella. However, many of these are of doubtful reliability because of the misidentification of
the parasitoid or its host (Robinson and Nielsen, 1993). Summaries are provided by Griswold
(1944), Hinton (1956) and Herting (1975). More recently, the endoparasitoid Apanteles carpatus
(Say) has been shown to be capable of completing development on T. bisselliella (Plarre et al.,
1999). This parasitoid shows a preference for older larval stages but can complete development on
all instars, and has been considered as a candidate for biological control of clothes moths in
households and museums (Plarre and Balnuweit, 2003). Zimmermann et al. (2003) tested 30
strains and species of the egg parasitoid Trichogramma, and the results of host acceptance and
preference tests indicated that T. piceum Dyurich was the most promising candidate for biological
control of T. bisselliella.
Nuclear polyhedrosis virus infections were first recorded in T. bisselliella by Lotmar (1941);
disease symptoms have been described by Whitfield and Cole (1958) and the histology and
ultrastructure by Hunter et al. (1973). The microsporidian Nosema has also been recorded as
attacking the larvae (Hinton, 1956). The bacterium Bacillus thuringiensis Berliner, has been shown
to kill T. bisselliella larvae when added to the larval diet (Yamvrias and Angus, 1969).
So far, there appear to have been no attempts made to use any of these organisms for the
biological control of clothes moths commercially.

5.6. Semiochemicals

Attempts to find alternatives to conventional pesticides for the control of T. bisselliella have
included investigating the use of antifeedants, antimetabolites that block insect metabolic reactions,
insect growth regulators, repellents and other chemicals controlling insect behaviour (Reagan, 1982).
For example, new antifeedants to control clothes moths in museum collections have been
investigated by Morita et al. (1988) and Zaitseva et al. (1990). Gerard et al. (1993) found
antifeedant properties against this species in extracts of leaves from a native New Zealand tree.
The use of antimetabolites for the control of clothes moths was examined by Pence (1959),
while Mauchamp (1974) studied the action of a juvenile hormone analogue on T. bisselliella. More
recently, Blattner et al. (1997) found high mortality when larvae were exposed to allosamidin
which inhibits chitinase and insect ecdysis.
Substances such as naphthalene, paradichlorobenzene and camphor have been used as
repellents in moth balls or moth flakes for many years (Chauvin and Vannier, 1994; Chio et al.,
1996). However, there are concerns about the hazards to humans of long-term exposure to these
chemicals and their use is being reviewed (Pinniger and Harmon, 1999). Insects are also repelled
by high doses of some traditional plant-derived materials including lavender, bay, rosemary and
wormwood. In recent years, plant products have been the subject of further investigations as safer
alternatives to conventional pesticides for clothes moth control (Gerard and Ruf, 1991; Gerard
et al., 1992; Detmers et al., 1992; Gerard and Ruf, 1995; Plarre et al., 1997; Blattner et al., 1997;
Ainge et al., 2002). However, it is worth remembering that some chemicals derived from plants are
as toxic as synthetic pesticides and the oils may have deleterious effects on some objects (Pinniger
and Winsor, 1998).
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The use of semiochemicals as novel control agents for stored-product insect pests has been
reviewed recently by Cox (2004). Approaches considered to have potential for improving the
protection of stored products included attractants for mass trapping, repellents as protective
bands and flushing agents, and the synchronised use of attractants and repellents to manoeuvre
pests away from infestable materials. It may be possible to use the pheromone traps described in
Section 5.1 of the current review to reduce levels of infestation through confusion or disruption of
the moth communication and mating system. Some of the semiochemicals referred to in Sections
3.2 and 3.4 could well be worth investigating further in this respect. In addition, semiochemical-
mediated location of host habitat by the parasitoid A. carpatus has been demonstrated by Takacs
et al. (1997), and it might be possible to use these chemicals to enhance the effectiveness of this
biocontrol agent for controlling infestations of T. bisselliella.

5.7. Modified atmospheric gases

Another method of insect control which has proved successful against T. bisselliella in some
situations is the use of modified atmospheric gases, particularly nitrogen and carbon dioxide, to
cause death by starving the insect of oxygen (Gilberg, 1991; Pinniger, 1991b; Koestler, 1992;
Selwitz and Maekawa, 1998). Although these gases are safer to use than traditional fumigants,
their main disadvantage is the length of time required to ensure control, particularly at lower
temperatures. The two main methods are by treatment with carbon dioxide at a concentration of
around 60% or by oxygen exclusion using nitrogen at concentrations of greater than 99.7% with
or without an oxygen scavenger (Pinniger, 2001). Bulky cylinders are usually required for the
nitrogen supply, although recently a method of on-site nitrogen generation has been developed
(Conyers, 2001). Both methods require the use of gas-proof sheeting or specially constructed gas-
tight chambers or polyethylene-aluminium laminate ‘bubbles’ to maintain conditions for up to 4
weeks (Pinniger, 1995; Newton et al., 1996).
One week in an atmosphere of 99.5% nitrogen at 30 1C was sufficient for complete kill of all life
stages of T. bisselliella, the egg being the most difficult stage to kill by anoxia (Gilberg, 1991; Rust
et al., 1996). In Japan, this moth is controlled by exposure at 25 1C to 60% carbon dioxide with
oxygen concentration at about 8% for 1 week (Kigawa et al., 2001). At an average temperature of
22 1C, a 2-week exposure was found to be necessary to ensure complete kill of T. bisselliella in
museum objects using similar gas concentrations in Canada (Warren, 2001).

5.8. Irradiation

Several different types of electromagnetic radiation have been examined for their abilities to kill
insects, including X-rays, beta and gamma rays, and microwaves (Nelson, 1966; Tilton, 1974;
Reagan, 1982; Pinniger, 1991a). In recent years gamma rays and microwaves have received most
attention. For example, Reyer (1992) reported good control of T. bisselliella infestations in
museum textiles using microwaves.
Irradiation as a means of disinfestation has the advantage of being faster than conventional
chemical methods and of leaving no harmful residues. However, microwaves can cause
undesirable side-effects through uneven heating, and they cannot be used on objects containing
metals. Gamma radiation units are expensive, require stringent safety precautions during
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operation, and can have deleterious effects on some materials such as plastics and glass (Pinniger
and Harmon, 1999).

6. Conclusions

Webbing clothes moths have been pests for at least 3000 years and it seems as if they will
continue to attack human clothing and other textiles for many years to come. Improvements in
storage coupled with targeted use of appropriate control measures could help to reduce
populations. However, the continuing problems with this pest highlight the need for more
effective methods of detection, prevention and management.

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