LICE (Phthiraptera)

(A) Ishnocera; (B)Anoplura; (C)Amblycera; (D) Rhynchopthirina.

Lice are a menace to humans, pets, and livestock, not only because of their blood-feeding or chewing habits, but also because of their ability to transmit pathogens. The human body louse has been indirectly responsible for influencing human history through its ability to transmit the causative agent of epidemic typhus. However, most of the 3200 known species of lice are ectoparasites of wild birds or mammals and have no known medical or veterinary importance. The order Phthiraptera is divided into two main taxonomic groups: the Anoplura (sucking lice) and Mallophaga (chewing or biting lice). All members of the Anoplura are obligate, hematophagous ectoparasites of placental mammals, whereas the more diverse.

Mallophaga include species that are obligate associates of birds, marsupials, and placental mammals. Although certain chewing lice imbibe blood, most species ingest host feathers, fur, skin, or skin products. Because of the different feeding strategies of the two groups, the blood-feeding Anoplura are far more important than the Mallophaga in transmitting pathogens to their hosts.

TAXONOMY

Major taxonomic syntheses for the sucldng lice include a series of eight volumes by Ferris (1919-1935) that remains the most comprehensive treatment of this group on a worldwide basis. Ferris (1951) updated much of his earlier work in a shorter overview of the group. Kim et al. (1986) have compiled an authoritative manual and identification guide for the sucking lice of North America. Durden and Musser (1994a) provide a taxonomic checldist for the sucking lice of the world, with host records and geographical distribution for each species. The chewing lice are taxonomically less well known than are the sucking lice, and few authoritative identification guides are available. These include a synopsis of the lice associated with laboratory animals (Kim et al. 1973), guides to the lice of domestic animals (Tuff 1977, Price and Graham 1997), and an identification guide to the lice of sub-Saharan Africa (Ledger 1980). These publications provide information on both sucking lice and chewing lice. Checklists of the Mallophaga of the world (Hopkins and Clay 1952) and of North America (Emerson 1972) are useful taxonomic references for this group.

Because of the relatively high degree of host specificity exhibited by both chewing and sucking lice, several host-parasite checklists have been prepared. These include a detailed list of both anopluran and mallophagan lice associated with mammals (Hopkins 1949), a hostparasite list for North American Mallophaga (Emerson 1972), a world host-parasite list for the chewing lice of mammals (Emerson and Price 1981 ), and a host-parasite checklist for the Anoplura of the world (Durden and Musser 1994b).

About 550 species of sucking lice have been described (Durden and Musser 1994a), all of which parasitize placental mammals; these lice are currently assigned to 50 genera and 15 families. About 2650 valid species of Mallophaga have been described; most of these are associated with birds, but about 400 (ca. 15%) parasitize mammals. The Mallophaga can be divided into 3 suborders (Table I), 11 families, and 205 genera.

The Mallophaga are divided into the following three groups (suborders of most authors): Amblycera (seven families, ca. 76 genera, and ca. 850 species), Ischnocera (three families, ca. 130 genera, and ca. 1800 species), and Rhyncophthirina (one family, 1 genus, and 3 species) (Figs. 4.1 and 4.5). However, there has been disagreement regarding the taxonomic rank of these three groups and their relationships to the Anoplura. Many current classifications treat the Phthiraptera as an order and assign suborder (or superfamily) rank to each of the Anoplura, Amblycera, Ischnocera, and Rhyncophthirina. Other classifications treat the Anoplura and Mallophaga as separate orders. Unfortunately, recent phylogenetic analyses of lice based on cladistic principles have produced contradictory results and have failed to resolve this issue. Regardless of current taxonomic interpretations, it is widely agreed that both sucking and chewing lice originated from a common nonparasitic ancestral group closely related to the order Psocoptera (book lice and bark lice). These two groups diverged in the late Jurassic or early Cretaceous Period, 100-150 million years ago.

Sucking lice of medical importance are assigned to two families, the Pediculidae and Pthiridae, whereas sucking lice of veterinary importance are assigned to five families: the Haematopinidae, Hoplopleuridae, Linognathidae, Pedicinidae, and Polyplacidae (Table II). Only one species of chewing louse, the dog biting louse, in the family Trichodectidae, has public health importance. Mallophaga of veterinary significance are typically placed in five families: the Boopiidae, Gyropidae, Menoponidae, Philopteridae, and Trichodectidae (Table I).

Classification and Hosts of Chewing Lice (Mallophaga) of Medical and Veterinary Importance

MORPHOLOGY

Lice are small (0.4-10 mm in the adult stage), wingless, dorso-ventrally flattened insects. The elongate abdomen possesses sclerotized dorsal, ventral, and/or lateral plates in many lice (Fig. 4.2); these provide some rigidity to the abdomen when it is distended by a blood meal or other food source. In adult lice the abdomen has 11 segments and terminates in genitalia and associated sclerotized plates. In females, the genitalia are accompanied by finger-like gonopods, which serve to guide, manipulate, and glue eggs onto host hair or feathers. The abdomen is adorned with numerous setae in most lice. Immature lice closely resemble adults but are smaller, have fewer setae,

(A) Ishnocera; (B)Anoplura; (C)Amblycera; (D) Rhynchopthirina.

FIGURE 4.1 Head and mouthparts ofrepresentatives of each ofthe four principal groups oflice. (A) Ishnocera; (B)Anoplura; (C)Amblycera; (D) Rhynchopthirina. (A, from Price and Graham, 1997; B and D, from Ferris, 1931; C, from Bedford, 1932)

and lack genitalia. After each nymphal molt, the abdomen is beset with progressively more setae, and the overall size of the louse increases.

The male genitalia in lice (Fig. 4.3) are relatively large and conspicuous, sometimes occupying almost half the length of the abdomen. The terminal, extrusable, sclerotized pseudopenis (aedeagus) is supported anteriorly by a basal apodeme. Laterally, it is bordered by a pair of chitinized parameres. Two or four testes are connected to the vas deferens, which coalesces posteriorly to form the vesicula seminalis. In the female, the vagina leads to a large uterus, to which several ovarioles supporting eggs in various stages of development are connected by the oviducts. Two or more large accessory glands, which secrete materials to coat the eggs, and a single spermatheca, in which sperm is stored, are situated posteriorly in the abdomen. Except for the human body louse, all lice cement their eggs, called nits, onto the hair or feathers of their host. Eggs are usually subcylindrical, with rounded ends and a terminal cap, the operculum (Fig. 4.4). On the top of the operculum is a patch of holes or areas with thin cuticle, called micropyles, through which the developing embryo respires. Most of the egg is heavily chitinized, which helps to protect the embryo from mechanical damage and desiccation. A suture of thin cuticle encircles the base of the operculum. At the time of hatching, the first-instar nymph emerges from the egg by cracldng this suture and pushing off the operculum.

In chewing lice, the head is broader than the thorax (Fig. 4.5). Amblyceran chewing lice have four-segmented antennae and have retained the maxillary palps characteristic of their psocopteran-like ancestor. However, ischnoceran chewing lice have three to five antennal segments and lack maxillary palps. In the Amblycera, the antennae are concealed in lateral grooves, whereas in the Ischnocera and Rhyncophthirina, the antennae are free from the head (Figs. 4.1 and 4.5).

There is a gradation in the specialization of the mouthparts and of the internal skeleton of the head, or tentorium, from the psocopteran-like ancestor of the lice through the Amblycera, Ischnocera, Rhyncophthirina, and Anoplura. Although mallophagan lice all possess chewing mouthparts (Fig. 4.6), the components and mechanics of these mouthparts differ for each group. In the Amblycera, the opposable mandibles move in a vertical plane, or perpendicular to the ventral surface of the head, whereas in the Ischnocera they move more or less horizontally. In contrast, the Rhyncophthirina possess tiny mandibles that are situated at the tip of an elongated rostrum (Figs. 4.1D and 4.5F). Through extreme

Classification and Hosts of Sucking Lice (Anoplura) of Medical and Veterinary Importance

modifications, members of the chewing louse genus Trochilocoetes (parasites of humming birds) have evolved mouthparts that can function as sucking organs.

The thorax in chewing lice usually appears as two, and occasionally three, segments. Chewing lice possess one or two simple claws on each leg; species that parasitize highly mobile hosts, especially birds, typically have two claws.

In sucking lice (Figs. 4.2 and 4.7) the head is slender and narrower than the thorax. Anoplura have three- to five-segmented antennae and lack maxillary palps. Eyes are reduced or absent in most sucking lice but are well developed in the medically important genera Pediculus and Pthirus (Fig. 4.7A, B), and ocular points, or eyeless projections posterior to the antennae, are characteristic of sucking lice in the genus Haematopinus (Fig. 4.7E).

As indicated by their name, anopluran mouthparts function as sucking devices during blood feeding

A generalized sucking louse (Anoplura), showing dorsal (left) and ventral (right) morphology.

FIGURE 4.2 A generalized sucking louse (Anoplura), showing dorsal (left) and ventral (right) morphology. (From Ignoffo, 1959.)

(Fig. 4.8). At rest, the mouthparts are withdrawn into the head and are protected by the snoutlike haustellum, representing the highly modified labrum. The haustellum is armed with tiny recurved teeth which hook into the host skin during feeding. The stylets, consisting of a serrated labium, the hypopharynx, and two maxillae, then puncture a small blood vessel (Fig. 4.8). The hypopharynx is a hollow tube through which saliva (containing anticoagulants and enzymes) is secreted. The maxillae oppose each other and are curved to form a food canal through which host blood is imbibed (Fig. 4.9).

In sucking lice, all three thoracic segments are fused and appear as one segment. In most species, the legs terminate in highly specialized claws for grasping the host

Internal abdominal anatomy of a male human body louse (Pediculus humanus humanus).

HGURE 4.3 Internal abdominal anatomy of a male human body louse (Pediculus humanus humanus). (From Ferris, 1951.)

Eggs (nits) of representative lice. (A) Chicken body louse, Menacanthus stramineus; (B) Oval guineapig louse, Gyropus ovalis; (C) Pigeon louse, Columbicola columbae; (D) Cattle biting louse, Bovicola bovis; (E) Elephant louse, Haematomyzus elephantis; (F) Human head louse, Pediculus humanus capitis. (From Marshall, 1981).

FIGURE 4.4 Eggs (nits) of representative lice. (A) Chicken body louse, Menacanthus stramineus; (B) Oval guineapig louse, Gyropus ovalis; (C) Pigeon louse, Columbicola columbae; (D) Cattle biting louse, Bovicola bovis; (E) Elephant louse, Haematomyzus elephantis; (F) Human head louse, Pediculus humanus capitis. (From Marshall, 1981).

pelage. These tibio-tarsal claws consist of a curved tarsal element which opposes a tibial spur (Fig. 4.10) to enclose a space that typically corresponds to the diameter of the host hair.

The internal anatomy of lice (Fig. 4.3) is best known for the human body louse. As in most hematophagous insects, strong cibarial and esophageal muscles produce a sucking action during blood feeding. The esophagus leads to a spacious midgut composed primarily of the ventriculus. The posterior region of the midgut is narrow and forms a connection between the ventriculus and the hindgut. Ventrally, mycetomes containing symbiotic microorganisms connect to the ventriculus.

LIFE HISTORY

Lice are hemimetabolous insects. Following the egg stage, there are three nymphal instars, the last of which molts to an adult. Although there is wide variation between species, the egg stage typically lasts for 4-15 days and each nymphal instar for 3-8 days; adults live for up to 35 days. Under optimal conditions many species of lice can complete 10-12 generations per year, but this is rarely achieved in nature. Host grooming, resistance, molting and feather loss, hibernation, and hormonal changes, as well as predators (especially insectivorous birds on large ungulates), parasites and parasitoids, and unfavorable weather conditions can reduce the number of louse generations.

Fecundities for fertilized female lice vary from 0.2 to 10 eggs per day. Males are unknown in some parthenogenetic species, whereas they typically constitute less than 5% of the adult population in the cattle biting louse and less than 1% in the horse biting louse.

BEHAVIOR AND ECOLOGY

Blood from the host is essential for the successful development and survival of all sucldng lice. Anoplura are vessel feeders, or solenophages, that imbibe blood through a hollow dorsal stylet derived from the hypopharynx (Fig. 4.9). Contraction of powerful cibarial and pharyngeal muscles create a sucking reaction for imbibing blood.

Chewing lice feed by the biting or scraping action of the mandibles. Bird-infesting chewing lice typically use their mandibles to sever small pieces of feather, which drop onto the labrum and are then forced into the mouth. Chewing lice which infest mammals use their mandibles in a similar manner to feed on host fur. Many chewing lice that infest birds and mammals can also feed on other integumental products, such as skin debris and secretions. Some species of chewing lice are obligate, or more frequently facultative, hematophages. Even those species of chewing lice that imbibe blood scrape the host integument until it bleeds. The rhyncophthirinan Haematomyzus elephantis, which parasitizes both African and Asian elephants, feeds in this manner.

Chewing lice (Mallophaga) of veterinary importance, showing dorsal morphology (left) and ventral morphology (right) in each case. Not drawn to scale. (A) Heterodoxus spiniger, male, from carnivores; (B) Tricholipeurus parallelus, female, from New World deer; (C) Sheepbiting louse (Bovicola bovis), female; (D) Dog-biting louse (Trichodectes canis), female; (E) Catbiting louse (Felicola subrostrata), male; (F) Elephant louse (Haematomyzus elephantis), male. (A-E, from Emerson and Price, 1975; F, from Werneck, 1950)

FIGURE 4.5 Chewing lice (Mallophaga) of veterinary importance, showing dorsal morphology (left) and ventral morphology (right) in each case. Not drawn to scale. (A) Heterodoxus spiniger, male, from carnivores; (B) Tricholipeurus parallelus, female, from New World deer; (C) Sheepbiting louse (Bovicola bovis), female; (D) Dog-biting louse (Trichodectes canis), female; (E) Catbiting louse (Felicola subrostrata), male; (F) Elephant louse (Haematomyzus elephantis), male. (A-E, from Emerson and Price, 1975; F, from Werneck, 1950)

Generalized mouthparts of an amblyceran chewing louse (Mallophaga). (A) Ventral view of head; (B) labium and associated structures; (C) mandibles. (Drawn by Margo Duncan)

FIGURE 4.6 Generalized mouthparts of an amblyceran chewing louse (Mallophaga). (A) Ventral view of head; (B) labium and associated structures; (C) mandibles. (Drawn by Margo Duncan)

Symbionts are thought to be present in all lice that imbibe blood. Symbionts in the mycetomes (Fig. 4.3) aid in blood meal digestion, and lice deprived of them die after a few days; female lice lacking symbionts also become sterile. In female human body lice, some symbionts migrate to the ovary, where they are transferred transovarially to the next generation of lice.

Lice in general exhibit host specificity, some to such a degree that they parasitize only one species of host. The hog louse, slender guineapig louse, large turkey louse, and several additional species listed in Tables I and II all are typical parasites of a single host species.

Host specificity is broader in some lice. Some lice of veterinary importance parasitize two or more closely related hosts. Examples include the three species which parasitize domestic dogs: Linognathus setosus, Trichodectes

Sucking lice (Anoplura) of medical and veterinary importance, showing dorsal morphology (left) and ventral morphology (right) in each case. Not drawn to scale. (A) Human body louse (Pediculus humanus humanus), female; (B) Human crab louse (Phthirus pubis), female; (C) Flying squirrel louse (Neohaematopinus sciuropteri), male; (D) Spirted rat louse (Polyplax spinulosa), male; (E) Hog louse (Haematopinus suis), female; (F) Little blue cattle louse (Solenoptes capillatus), male; (G) Dog sucking louse (Linognathus setosus), male; (H) Longnosed cattle louse (L. vituli), female. (From Ferris, 1923-1935.)

FIGURE 4.7 Sucking lice (Anoplura) of medical and veterinary importance, showing dorsal morphology (left) and ventral morphology (right) in each case. Not drawn to scale. (A) Human body louse (Pediculus humanus humanus), female; (B) Human crab louse (Phthirus pubis), female; (C) Flying squirrel louse (Neohaematopinus sciuropteri), male; (D) Spirted rat louse (Polyplax spinulosa), male; (E) Hog louse (Haematopinus suis), female; (F) Little blue cattle louse (Solenoptes capillatus), male; (G) Dog sucking louse (Linognathus setosus), male; (H) Longnosed cattle louse (L. vituli), female. (From Ferris, 1923-1935.)

Head region of a sucking louse (Anoplura) feeding on a host, showing components of mouthparts and associated internal structures. (Original by Margo Duncan.)

FIGURE 4.8 Head region of a sucking louse (Anoplura) feeding on a host, showing components of mouthparts and associated internal structures. (Original by Margo Duncan.)

Cross-section through the mouthparts of a sucking louse (Anoplura). (Original by Margo Duncan.)

FIGURE 4.9 Cross-section through the mouthparts of a sucking louse (Anoplura). (Original by Margo Duncan.)

canis, and Heterodoxus spiniger. These lice also parasitize foxes, wolves, coyotes, and occasionally other carnivores. Similarly, the horse sucking louse (Haematopinus asini), parasitizes horses, donkeys, asses, mules, and zebras, whereas L. africanus parasitizes both sheep and goats. At least six species of chewing lice are found on domestic fowl, all of them parasitizing chickens, but some also feeding on turkeys, guinea fowl, pea fowl, or pheasants (Table I). Lice found on atypical hosts are termed stragglers.

Some sucking lice, such as the three taxa that parasitize humans, the sheep foot louse, and the sheep face louse, are not only host specific, but also infest specific body areas, from which they can spread in severe infestations. Many chewing lice, particularly species that parasitize birds, also exhibit both host specificity and site specificity; examples include several species that are found on domestic fowl, and species confined to turkeys, geese, and ducks (Table I). Lice inhabiting different body regions on the same host typically have evolved morphological adaptations in response to specific attributes of the host site. These include characteristics such as morphological differences of the pelage, thickness of the skin, availability of blood vessels, and grooming or preening activities of the host. Site specificity in chewing lice is most prevalent in the more sedentary, specialized Ischnocera than in the mostly mobile, morphologically unspecialized Amblycera. For example, on many bird hosts, roundbodied ischnocerans with large heads and mandibles are predominately found on the head and neck. Elongate forms with narrow heads and small mandibles tend to inhabit the wing feathers, whereas morphologically intermediate forms occur on the back and other parts of the body.

Some chewing lice inhabit highly specialized host sites. These include members of the amblyceran genus Piagetiella, which are found inside the oral pouches of pelicans, and members of several amblyceran genera, including Actornithophilus and Colpocephalum, which live inside feather quills. Several bird species are parasitized by 5 or more different species of site-specific chewing lice, and up to 12 species may be found on the neotropical bird Crypturellus soul (a tinamou).

Site specificity is less well documented for sucking lice. However, domestic cattle may be parasitized by as many as five anopluran species, each predominating on particular parts of the body. Similarly, some Old World squirrels and rats can support up to six species of sucking lice.

Because of the importance of maintaining a permanent or close association with the host, lice have evolved specialized host-attachment mechanisms to resist grooming activities of the host. The robust tibio-tarsal claws of sucking lice (Fig. 4.10) are very important in securing them to their hosts. Various arrangements of hooks and spines, especially on the heads of lice that parasitize arboreal or flying hosts, such as squirrels and birds, also aid in host attachment. Mandibles are important attachment appendages in ischnoceran and rhyncophthirinan chewing lice. In some species of Bovicola, a notch in the first antenhal segment encircles a host hair to facilitate attachment.

Tibio-tarsal claws and antenna of Linognathus africanis (Anoplura): scanning electron micrograph.

FIGURE 4.10 Tibio-tarsal claws and antenna of Linognathus africanis (Anoplura): scanning electron micrograph. (From Price and Graham, 1997.)

A few lice even possess ctenidia (“combs”) that are convergently similar in morphology to those characteristic of many fleas. They occur most notably among lice that parasitize coarse-furred, arboreal, or flying hosts. Additionally, chewing lice that parasitize arboreal or flying hosts often have larger, more robust claws than do their counterparts that parasitize terrestrial hosts.

Because of their reliance on host availability, lice are subjected to special problems with respect to their longterm survival. MI sucking lice are obligate blood-feeders; even a few hours away from the host can prove fatal to some species. Some chewing lice also are hematophages and similarly cannot survive prolonged periods off the host. However, many chewing lice, particularly those that subsist on feathers, fur, or other skin products, can survive for several days away from the host. For example, the cattle biting louse can survive for up to 11 days (this species will feed on host skin scrapings), and Menacanthusspp. of poultry can survive for up to 3 days off the host. Off-host survival is generally greater at low temperatures and high humidities. At 26~ and 65% relative humidity (RH), 4% of human body lice die within 24 hr, 20% within 40 hr, and 84% within 48 hr. At 75% RH, a small proportion of sheep foot lice survives for 17 days at 2~ whereas most die within 7 days at 22~ Recently fed lice generally survive longer than unfed lice away from the host. Mthough most lice are morphologically adapted for host attachment and are disadvantaged when dislodged, the generalist nature of some amblyceran chewing lice better equips them for locating another host by crawling across the substrate. Amblycerans are more likely than other lice to be encountered away from the host, accounting for observations of these lice on bird eggs or in unoccupied nests and roosts.

Host grooming is an important cause of louse mortality. Laboratory mice infested by the mouse louse, for example, usually limit their louse populations to 10 or fewer individuals per mouse by regular grooming. Prevention of self-grooming or mutual grooming by impaired preening action of the teeth or limbs of such mice can result in heavy infestations of more than 100 lice. Similarly, impaired preening due to beak injuries in birds can result in tremendous increases of louse populations. Biting, scratching, and licking also reduce louse populations on several domestic animals.

Whereas most species of lice on small and mediumsized mammals exhibit only minor seasonal differences in population levels, some lice associated with larger animals show clear seasonal trends. Some of these population changes have been attributed to host molting, fur density and length, hormone levels in the blood meal, or climatological factors such as intense summer heat, sunlight, or desiccation. On domestic ungulates in temperate regions, louse populations typically peak during the winter or early spring and decline during the summer. An exception to this trend is the cattle tail louse, whose populations peak during the summer.

Another important aspect of louse behavior is the mode of transfer between hosts. Direct host contact appears to be the primary mechanism for louse exchange. Transfer of lice from an infested mother to her offspring during suclding (in mammals) or during nest sharing (in birds and mammals) is an important mode of transfer. Several species of lice that parasitize livestock transfer during suclding, including the sheep face louse and the sheep biting louse, both of which move from infested ewes to their lambs at this time. Lice can also transfer during other forms of physical contact between hosts, such as mating or fighting. Transfer of lice between hosts also can occur between hosts that are not in contact. The sheep foot louse, for example, can survive for several days off the host and reach a new host by crawling across pasture land. Nests of birds and mammals can act as foci for louse transfer, but these are infrequent sites of transfer.

Dispersal of some lice occurs via phoresy, in which they temporarily attach to other arthropods and are carried from one host to another (Fig. 4.11). During phoresy, most lice attach to larger, more mobile bloodfeeding arthropods, usually a fly, such as a hippoboscid or muscoid. Phoresy is particularly common among ischnoceran chewing lice. Movement of the mouthparts in a horizontal plane better facilitates their attachment to a fly than in the amblycerans, in which mouthparts move in a vertical plane. Phoresy is rare among sucking lice. This is

Two ischnoceran chewing lice (Mallophaga) phoretic on a hippoboscid fly, attached by their mandibles to the posterior abdomen.

FIGURE 4.11 Two ischnoceran chewing lice (Mallophaga) phoretic on a hippoboscid fly, attached by their mandibles to the posterior abdomen. (From Rothschild and Clay, 1952).

probably because attachment to the fly is achieved by the less efficient mechanism of grasping with the tarsal claws.

Mating in lice occurs on the host. It is initiated by the male pushing his body beneath that of the female and curling the tip of his abdomen upward. In the human body louse, the male and female assume a vertical orientation along a hair shaft, with the female supporting the weight of the male as he grasps her with his anterior claws. Most lice appear to exhibit similar orientation behavior during mating. Notable exceptions include the crab louse of humans, in which both sexes continue to clasp with their claws a host hair, rather than each other, during mating; and the hog louse, in which the male strokes the head of the female during copulation. Some male ischnoceran chewing lice possess modified hooklike antennal segments, with which they grasp the female during copulation.

Oviposition behavior by female lice involves crawling to the base of a host hair or feather and cementing one egg at a time close to the skin surface. Two pairs of fingerlike gonopods direct the egg into a precise location and orientation as a cement substance is secreted around the egg and hair base. Optimal temperature requirements for developing louse embryos inside eggs are very narrow, usually within a fraction of a degree, such as may occur on a precise area on the host body. For this reason, female lice typically oviposit preferentially on an area of the host that meets these requirements.

LICE OF MEDICAL INTEREST

Three taxa of sucking lice parasitize humans throughout the world: the body louse, head louse, and crab louse (pubic louse). All are specific ectoparasites of humans; rarely, dogs or other companion animals may have temporary, selflimiting infestations.

Human head and body lice are closely related and can interbreed to produce fertile offspring in the laboratory. For this reason, they generally are recognized as separate subspecies of Pediculus humanus, as in this chapter. Nevertheless, they rarely interbreed in nature, which has prompted some epidemiologists to treat them as separate species, P. humanus (body louse) and P. capitis (head louse).

Human body louse (Pediculus humanus humanus)

The human body louse (Figs. 4.7A and 4.12) or cootie was once an almost ubiquitous companion of humans. Today it is less common, especially in developed nations. Body lice persist as a significant problem in less developed nations in parts of Africa, Asia, and Central and South America. This is significant because P.h. humanus is the only louse of humans that is known

FIGURE 4.12 Human body lice (Pediculus humanus humanus) feeding on a human. (Courtesy of Elton J. Hansens.)

FIGURE 4.12 Human body lice (Pediculus humanus humanus) feeding on a human. (Courtesy of Elton J. Hansens.)

to naturally transmit pathogens. The large-scale reduction in body louse infestations worldwide has led to a concomitant decrease in the prevalence of human louse-borne diseases. However, situations that result in human overcrowding and unsanitary conditions (e.g., wars, famines, and natural disasters) can lead to a resurgence of body louse infestations, often accompanied by one or more louse-borne diseases.

Adult human body lice (Figs. 4.7A and 4.12) are 2.3-3.6 mm long. Under optimal conditions their populations can multiply dramatically if unchecked; e.g., if clothes of infested individuals are not changed and washed in hot water at regular intervals. In unusually severe infestations, populations of more than 30,000 body lice on one person have been recorded. Body lice typically infest articles of clothing and crawl onto the body only to feed. Females lay an average of four or five eggs per day, and these typically hatch after 8 days. Unique among lice, females oviposit not on hair, but on clothing (Fig. 4.13), especially along seams

FIGURE 4.13 Eggs of the human body louse (Pediculus humanus humanus) attached to clothing. (Courtesy ofElton J. Hansens.)

FIGURE 4.13 Eggs of the human body louse (Pediculus humanus humanus) attached to clothing. (Courtesy ofElton J. Hansens.)ฃ

and creases. Each nymphal instar lasts for 3-5 days, and adults can live for up to 30 days.

Biting by body lice often causes intense irritation, with each bite site typically developing into a small red papule with a tiny central clot. The bites usually itch for several days but occasionally for a week or more. Persons exposed to numerous bites over long periods often become desensitized and show little or no reaction to subsequent bites. Persons with chronic body louse infestations may develop a generalized skin thickening and discoloration called Vagabond disease or Hobo disease, names depicting a lifestyle that can promote infestation by body lice. Several additional symptoms may accompany chronic infestations. These include lymphadenopathy (swollen lymph nodes), edema, increased body temperature often accompanied by fever, a diffuse rash, headache, joint pain, and muscle stiffness.

Some people develop allergies to body lice. Occasionally, patients experience a generalized dermatitis in response to one bite or small numbers of bites. A form of asthmatic bronchitis has similarly been recorded in response to allergy to louse infestations. Secondary infections such as impetigo or blood poisoning can also result from body louse infestations.

Body lice tend to leave persons with elevated body temperatures and may crawl across the substrate to infest a nearby person. This has epidemiological significance because high body temperatures of lousy persons often result from fever caused by infection with louseborne pathogens.

Human head louse (Pediculus humanus cap#is)

The human head louse is virtually indistinguishable from the human body louse on the basis of morphological characters and its life cycle. Unless a series of specimens is available for analysis it is often impossible to separate the two subspecies. Generally, adult head lice are slightly smaller (2.1-3.3 mm in length) than body lice.

As indicated by their name, human head lice typically infest the scalp and head region, rather than other areas of the body infested by body lice. Females attach their eggs to the base of individual hairs. As the hair grows, the eggs become further displaced from the scalp. An indication of how long a patient has been infested can be gleaned by measuring the farthest distance of eggs from the scalp and comparing this to the growth rate of hair.

Today, head lice are far more frequently encountered than body lice, especially in developed countries. Transmission occurs by person-to-person contact and via shared objects such as combs, brushes, headphones, and caps. School-age children are at high risk because they are often more likely to share such items. Some school districts in the United States and Britain have infestation prevalences approaching 50% in students. It has been estimated that 6-12 million people, principally children, are infested with head lice annually in the United States. Some ethnic groups, such as persons of African origin, have coarser head hairs and are less prone to head louse infestations. The reason for this is simply that the tibio-tarsal claws of these lice cannot efficiently grip the thicker hairs.

Although head lice are not known to transmit pathogens, heavy infestations can cause severe irritation. As is the case with human body lice, the resultant scratching often leads to secondary infections such as impetigo, pyoderma, or blood poisoning. Severe head louse infestations occasionally result in the formation of scabby crusts beneath which the lice tend to aggregate. Enlarged lymph nodes in the neck region may accompany such infestations.

Human crab louse (Pthirus pubis)

The crab louse, or pubic louse, is a medium-sized (1.1- 1.8 mm long), squat louse (Fig. 4.7B), with robust tibio-tarsal claws used for grasping thick hairs, especially those in the pubic region. It also may infest coarse hairs on other parts of the body, such as the eyebrows, eyelashes, chest hairs, beards, moustaches, and armpits. This louse typically transfers between human partners during sexual intercourse and other intimate contact; in France, crab lice are described as “papillons d’amour” (butterflies of love). Transfer via infested bed linen or toilet seats can also occur. This is uncommon, however, because crab lice can survive for only a few hours off the host.

Female crab lice lay an average of three eggs per day. Eggs hatch after 7-8 days; the three nymphal instars together last for 13-17 days. Under optimal conditions the generation time is 20-25 days. The intense itching caused by these lice is often accompanied by purplish lesions at bite sites and by small blood spots from squashed lice or louse feces on underwear. Crab lice are widely distributed and relatively common throughout the world. They are not known to transmit any pathogens. One epidemiological study, however, revealed a positive relationship between infection with hepatitis B virus and crab louse infestation.

LICE OF VETERINARY INTEREST

A wide variety of lice infests domestic, livestock, and laboratory animals (Tables I and II). Many hosts, particularly small rodents, often support few if any lice, whereas large hosts such as livestock animals, including poultry, may be parasitized by very large numbers of lice. For example, fewer than 10 mouse lice (Polyplax serrata) on a house mouse are a typical burden, but more than a million lice may be present on extremely heavily infested sheep, cattle, horses, or other large animals.

LICE OF CATTLE

Lice are a major problem in cattle operations worldwide. Domestic cattle are parasitized by six species of lice: three species of Haematopinus, one of Linognathus, one of Solenopotes, and one of Bovicola. Domestic Asiatic buffalo are typically parasitized by H. tuberculatus (Tables I and II).

Females of the cosmopolitan cattle biting louse (Bovicola bovis) lay an average of 0.7 eggs per day, which hatch 7-10 days later. Each nymphal instar lasts 5-6 days, and adult longevity can be as long as 10 weeks. Preferred host sites for this louse are the base of the tail, the shoulders, and the top line of the back, but lice may also populate the pollard in severe infestations.

The longnosed cattle louse (L. vituli) (Fig. 4.7H) also is a worldwide pest. Females deposit about one egg per day, and the life cycle is completed in about 21 days. This louse is most common on calves and dairy stock; it rarely occurs in large numbers on mature cattle. Preferred infestation sites are the dewlap and shoulders; declining spring populations are often confined to the shoulders.

The little blue cattle louse (Solenopotes capillatus) (Fig. 4.7F) also has a worldwide distribution. Females lay one or two eggs per day; oviposition typically causes the hairs on which eggs are laid to bend. Eggs hatch after about 10 days, and adulthood is reached about 11 days later. Clusters of S. capillatus typically occur on the muzzle, dewlap, and neck of mature cattle. Aggregations of this louse may surround the eyes in severely infested animals, giving a spectacled appearance to the host.

The cosmopolitan shortnosed cattle louse ( H. eurysternus) is the largest louse found on North American cattle; adults measure 3.5-4.7 mm in length. Females lay one to four eggs per day for about 2 weeks, nymphs reach adulthood in about 14 days, and adult longevity is 10-15 days. This louse is more common on mature cattle than on young animals. Preferred infestation sites are the top of the neck, the dewlap, and brisket. However, in severe infestations, the entire region from the base of the horns to the base of the tail can be infested. In North America, H. eurysternus is most prevalent in the Great Plains and Rocky Mountain regions.

The cattle tail louse ( H. quadripertusus) parasitizes cattle in the warmer regions of the world. It was inadvertently introduced into the United States, where it now occurs in the Gulf Coast states. Females of this louse oviposit on the tail hairs, which become matted with eggs in severe infestations. Infested tail heads may be shed under these circumstances. Eggs hatch after 9-25 days, depending on the season. Under optimal conditions, the entire life cycle can be as short as 25 days. Nymphs migrate over the host body surface, but adults are typically confined to the tail head. Unlike other cattle lice, H. quadripertusus is most abundant during the summer.

Except for H. quadripertusus, cattle lice increase in numbers during the winter and early spring in temperate regions. During summer, lice persist on 1-2% of the members of a herd; these chronically infested animals typically reinfest other herd members during the winter. Bulls and older cows often serve as reservoirs of lice. Bulls have longer, thicker hair and massive shoulders and neck that compromise self-grooming. During summer, a small number of lice can survive on the cooler ear tips, where lethal temperatures are rarely reached.

LICE OF OTHER LIVESTOCK ANIMALS

Horses, donkeys, hogs, goats, and sheep are parasitized by one or more species of louse (Tables I and II). Except for hogs, all of these animals are parasitized by both sucking lice and chewing lice. The horse biting louse (B. equi) is the most important louse of equids worldwide. Females of this louse oviposit on fine hairs, avoiding the coarse hairs of the mane and tail. This louse typically infests the side of the neck, the flanks, and tail base but can infest most of the body (except the mane, tail, ears, and lower legs) in severe infestations. Longhaired horse breeds are more prone to infestation by B. equi.

Domestic swine are parasitized by the hog louse (H. suis) (Fig. 4.7E). This is a large species in which adult females measure ca. 5 mm in length. Hog lice usually frequent skin folds of the head (especially the ears), neck, shoulders, and flanks of swine. Female hog lice lay an average of 3.6 eggs per day. These are deposited singly on hairs along the lower parts of the body, in skin folds on the neck, and on and in the ears. Eggs typically hatch 13-15 days later; each nymphal instar lasts 4-6 days. Adult hog louse longevity can be up to 40 days, and 6-15 generations can be completed per year, depending on environmental conditions.

Domestic sheep and goats are parasitized by several species of sucking lice and chewing lice (Tables I and II). One of these, L. africanus, parasitizes both hosts. Lice of sheep and goats, especially chewing lice, are economically important wherever these livestock animals are farmed, but especially in Australia, New Zealand, and the United States. Females of the sheep biting louse (B. ovis) lay one or two eggs per day and can live for up to 30 days; each nymphal instar lasts 5-9 days. B. ovis mainly infests the back and upper parts of the body but may populate the entire body in severe infestations. This louse causes intense irritation, and infested sheep typically rub against fences and trees, tearing the fleece and greatly reducing its value. Sucldng louse infestations of sheep rarely cause major economic problems.

LICE OF CATS AND DOGS

Domestic cats are parasitized by one species of chewing louse, whereas dogs are parasitized by two species of chewing lice and one species of sucking louse. All four species seem to be distributed worldwide, but none is a common associate of healthy cats or dogs in North America.

The cat biting louse (Felicola subrostrata) (Fig. 4.5E) parasitizes both domestic and wild cats. It may occur almost anywhere on the body. Both the dog biting louse ( T. canis) (Fig. 4.5D) and the dog sucking louse (L. setosus) (Fig. 4.7G) parasitize dogs and closely related wild canids. For example, T. canis also parasitizes coyotes, foxes, and wolves. A second species of chewing louse of dogs is Heterodoxus spiniger (Fig. 4.5A), which evolved in Australasia from marsupial-infesting lice and apparently switched to dingo hosts. It now parasitizes various canids and other carnivores throughout the world. T. can# usually infests the head, neck, and tail region of dogs, where it attaches to the base of a hair using its claws or mandibles. L. setosus occurs primarily on the head and neck and may be especially common beneath collars. H. spiniger can typically be found anywhere on its host.

LICE OF LABORATORY ANIMALS

The principal species of lice that parasitize laboratory mammals have been described by Kim et al. (1973). These lice also parasitize feral populations of their respective hosts. The house mouse (Mus musculus) is often parasitized by the mouse louse (P. serrata). Populations of this louse are typically low, with 10 or fewer lice per infested mouse, unless self-grooming or mutual host grooming is compromised. Eggs of this louse typically hatch 7 days after oviposition. Together the three nymphal instars last only 6 days under optimal conditions, which can result in a generation time as short as 13 days. Domestic rats are often parasitized by the spined rat louse (P. spinulosa) (Fig. 4.7D) and the tropical rat louse (Hoplopleura pacifica). Common hosts include the black rat (Rattus rattus) and the Norway rat (R. norvegicus). The spined rat louse parasitizes these hosts throughout the world, whereas the tropical rat louse is confined to tropical, subtropical, or warm temperate regions, including the southern United States.

Laboratory rabbits are parasitized by the rabbit louse (Haemodipsus ventricosis). This louse originated in Europe but has accompanied its host and been introduced throughout the world.

LICE OF POULTRY AND OTHER BIRDS

At least nine species of chewing lice commonly infest poultry (Table I) in various parts of the world. Individual birds can be parasitized by multiple species, each of which often occupies a preferred host site. The chicken body louse (Menacanthus stramineus) (Fig. 4.14) is the most common and destructive louse of domestic chickens. It has a worldwide distribution and often reaches pest proportions. Adults measure 3-3.5 mm in length. Females lay one or two eggs per day, cementing them in clusters at the bases of feathers, especially around the vent. Eggs typically hatch after 4-5 days. Each nymphal instar lasts about 3 days, and the generation time typically is 13-14 days. These lice are most abundant on the sparsely feathered vent, breast, and thigh regions. Several other chewing lice are pests of poultry more or less throughout the world (Table I). Adults of the shaft louse (Menopon gallinae) measure ca. 2 mm in length, and females deposit eggs singly at the base of the shaft on thigh and breast feathers. Eggs of the wing louse (Lipeurus capon#) hatch 4-7 days after the female has cemented them to the base of a feather. Nymphal stages of this species each last 5-18 days; generation time typically is 18-27 days, and females can live up to 36 days. Females of the chicken head louse (Cuclutogaster heterographus) attach their eggs to the bases of downy feathers. Eggs hatch after 5-7 days, each nymphal instar lasts 6-14 days, and average generation time is 35 days.

FIGURE 4.14 Chicken body lice (Menacanthus stramineus) on a chicken. (Courtesy of Nancy C. Hinlde.)

FIGURE 4.14 Chicken body lice (Menacanthus stramineus) on a chicken. (Courtesy of Nancy C. Hinlde.)

Poultry lice typically transfer to new birds by direct host contact. However, because most species can survive for several hours or days off the host, they also can infest new hosts during transportation in inadequately disinfected cages or vehicles.

PUBLIC HEALTH IMPORTANCE

Three important pathogens are transmitted to humans by body lice. These are the agents of epidemic typhus, trench fever, and louse-borne relapsing fever. Today, the prevalence and importance of all three of these louse-borne diseases are low compared to times when human body lice were an integral part of human life. However, trench fever has emerged as an opportunistic disease of immunocompromised individuals, including persons who are positive for human immunodeficiency virus (HIV).

EPIDEMIC TYPHUS

Epidemic typhus is a rickettsial disease caused by infection with Rickettsia prowazekii. It is also known as louseborne fever, jail fever, and exanthematic typhus. The disease persists in several parts of the world, most notably in Burundi, Democratic Republic of Congo, Ethiopia, Nigeria, Rwanda, and areas of northeastern and central Africa, Russia, Central and South America, and northern China. Epidemic typhus is largely a disease of cool climates, including higher elevations in the tropics. It thrives in conditions of widespread body louse infestations, overcrowding, and poor sanitary conditions. Epidemic typhus apparently was absent from the New World until the 1500s, when the Spanish introduced the disease. One resulting epidemic in 1576-1577 killed 2 million Indians in the Mexican highlands alone. The vector of R. prowazekii is the human body louse. Lice become infected when they feed on a person with circulating /L prowazekii in the blood. Infective rickettsiae invade cells that line the louse gut and multiply there, eventually causing the cells to rupture. Liberated rickettsiae either reinvade gut cells or are voided in the louse feces. Other louse tissues typically do not become infected. Because salivary glands and ovaries are not invaded, anterior-station and transovarial transmission do not occur. Infection of susceptible humans occurs via louse feces (posterior-station transmission) when infectious rickettsiae are scratched into the skin in response to louse bites. R. prowazekii can remain viable in dried louse feces for 60 days. Infection by inhalation of dried louse feces or by crushed lice are less frequent means of contracting the disease.

Transmission of R. prowazekii by body lice was first demonstrated by Charles Nicolle, working at the Institut Pasteur in Tunis in 1909. During these studies, Nicolle accidentally became infected with epidemic typhus, from which he fortunately recovered. He was awarded the Nobel prize in 1928 for his groundbreaking work on typhus. Several other typhus workers also were infected with R. prowazekii during laboratory experiments. The American researcher Howard T. Ricketts, working in Mexico, and Czech scientist Stanislaus yon Prowazek, working in Europe, both died from their infections and were recognized posthumously when the etiologic agent was named. Infection with R. prowazekii is ultimately fatal to body lice as progressively more and more infected gut cells are ruptured. Infective rickettsiae are first excreted in louse feces 3-5 days after the infective blood meal. Lice usually succumb to infection 7-14 days after the infectious blood meal, although some may survive to 20 days. The disease caused by infection with R. prowazekii and transmitted by body lice is called classic epidemic typhus because it was the first form of the disease to be recognized. Disease onset occurs relatively soon after infection by a body louse in classic epidemic typhus. Symptoms generally appear after an incubation period of 10-14 days. Abrupt onset of fever, accompanied by malaise, muscle and head aches, cough, and general weakness, usually occurs at this time. A blotchy rash spreads from the abdomen to the chest and then often across most of the body, typically within 4-7 days following the initial symptoms. The rash rarely spreads to the face, palms, and soles, and then only in severe cases. Headache, rash, prostration, and delirium intensify as the infection progresses. Coma and very low blood pressure often signal fatal cases. A case fatality rate of 10-20% is characteristic of most untreated epidemics, although figures approaching 50% have been recorded. Diagnosis of epidemic typhus involves the demonstration of positive serology, usually by microimmunofluorescence. DNA primers specific to R. prowazekii can also be amplified by polymerase chain reaction from infected persons or lice. One-time antibreak biotic treatment, especially with doxycycline, tetracycline, or chloramphenicol, usually results in rapid and complete recovery. Vaccines are available but are not considered to be sufficiently effective for widespread HSC. Persons that recover from epidemic typhus typically harbor R. prowazekii in lymph nodes or other tissues for months or years. This enables the pathogen to again invade other body tissues to cause disease seemingly at any time. This form of the disease is called recrudescent typhus or Brill-Zinsser disease. The latter name recognizes two pioneers in the study of epidemic typhus: Nathan Brill, who first recognized and described recrudescent typhus in 1910, and Hans Zinsser, who demonstrated in 1934 that it is a form of epidemic typhus. Zinsser’s (1935) book Rats, Lice, and History is a pioneering account of the study of epidemic typhus in general.

Recrudescent typhus was widespread during the 19th and early 20th centuries in some of the larger cities along the east coast of the United States (e.g., Boston, New York, and Philadelphia). At that time, immigrants from regions that were rampant with epidemic typhus, such as eastern Europe, presented with Brill-Zinsser disease after being infected initially in their country of origin. Some of these patients experienced relapses more than 30 years after their initial exposure, with no overt signs of infection with R. prowazekii between the two disease episodes. Because infestation with body lice was still a relatively common occurrence during that period, the lice further disseminated the infection to other humans, causing local outbreaks. The last outbreak of epidemic typhus in North America occurred in Philadelphia in 1877. Today, even recrudescent typhus is a rare occurrence in North America. However, this form of typhus is still common in parts of Africa, Asia, South America, and, occasionally, in eastern Europe. The southern flying squirrel ( Glaucomys volans) has been identified as a reservoir of R. prowazekii in the United States, where it has been found to be infected in Virginia during vertebrate serosurveys for Rocky Mountain spotted fever. Since the initial isolations from flying squirrels in 1963, R. prowazekii has been recorded in flying squirrels and their ectoparasites in several states, especially eastern and southern states. Peak seroprevalence (about 90%) in the squirrels occurs during late autumn and winter, when fleas and sucldng lice are also most abundant on these hosts. Although several ectoparasites can imbibe R. prowazekii when feeding on infected flying squirrels, only the sucldng louse Neohaematopinus sciuropteri (Fig. 4.7C) is known to maintain the infection and transmit the pathogen to uninfected squirrels. Several cases of human infection have been documented in which the patients recalled having contact with flying squirrels, especially during the winter months when these rodents commonly occupy attics of houses. To distinguish this form of the disease from classic and recrudescent typhus, it is called sporadic epidemic typhus or sylvatic epidemic typhus. Many details, such as the prevalence and mode of human infection, remain unresolved. Because the louse N. sciuropteri does not feed on humans, it is speculated that human disease occurs when infectious, aerosolized particles of infected louse feces are inhaled from attics or other sites occupied by infected flying squirrels. Except for flying squirrels in North America, humans are the only proven reservoirs of R. prowazekii. Widespread reports published in the 1950s to 1970s that various species of ticks and livestock animals harbored R. prowazekii have since been disproved. Historically, epidemic typhus has been the most widespread and devastating of the louse-borne diseases. Zinsser (1935) and Snyder (1966) have documented the history of this disease and highlighted how major epidemics have influenced human history. For example, the great outbreak of disease at Athens in 430 BC, which significantly influenced the course of Greek history, appears to have been caused by epidemic typhus. Napoleon’s vast army of 1812 was defeated more by epidemic typhus than by opposing Russian forces. Soon thereafter (ca. 1816- 1819), 700,000 cases of epidemic typhus occurred in Ireland. Combined with the potato famine of that period, this encouraged many people to emigrate to North America; some of these people carried infected lice or latent infections with them. During World War II, several military operations in North Africa and the Mediterranean region were hampered by outbreaks of epidemic typhus. One epidemic in Naples in 1943 resuited in over 1400 cases and 200 deaths. This outbreak is particularly noteworthy because it was the first epidemic of the disease to be interrupted by human intervention through widespread application of the insecticide dichlorodiphenyltrichloroethane (DDT) to louseinfested persons. Today, epidemic typhus is much less of a health threat than it once was. This is largely because few people, especially in developed countries, are currently infested by body lice. Higher sanitary standards, less overcrowding, regular laundering and frequent changes of clothes, effective pesticides, and medical advances have contributed to the demise of this disease. Nevertheless, epidemic typhus has the potential to re-emerge. This is evidenced by the largest outbreak of epidemic typhus since World War II that affected about half a million people living in refugee camps in Burundi in 1997-1998. Similarly, more than 5600 cases were recorded in China during 1999. Additional information about epidemic typhus is provided by the Pan American Health Organization/World Health Organization (1973), McDade (1987), and Azad (1988).

LOUSE-BORNE RELAPSING FEVER

Also known as epidemic relapsing fever, this disease is caused by the spirochete bacterium Borrelia recurrentis. This pathogen is transmitted to humans by the human body louse, as first demonstrated by Sergent and Foley in 1910. Clinical symptoms include the sudden onset of fever, headache, muscle ache, anorexia, dizziness, nausea, coughing, and vomiting. Thrombocytopenia (a decrease in blood platelets) also can occur and cause bleeding, which may initially be confused for a symptom of a hemorrhagic fever. Episodes of fever last 2-12 days (average, 4 days), typically followed by periods of 2-8 days (average, 4 days) without fever, with two to five relapses being usual. As the disease progresses, the liver and spleen enlarge rapidly, leading to abdominal discomfort and labored, painful breathing as the lungs and diaphragm are compressed. At this stage, most patients remain quietly prostrate with a glazed expression, often shivering and taking shallow breaths. Mortality rates for untreated outbreaks range from 5 to 40%. Antibiotic treatment is with penicillin or tetracycline. Humans are the sole known reservoir of B. recurrentis. Body lice become infected when they feed on an infected person with circulating spirochetes. Most of the spirochetes perish when they reach the louse gut, but a few survive to penetrate the gut wall, where they multiply to massive populations in the louse hemolymph, nerves, and muscle tissue. Spirochetes do not invade the salivary glands or ovarian tissues and are not voided in louse feces. Therefore, transmission to humans occurs only when infected lice are crushed during scratching, which allows the spirochetes in infectious hemolymph to invade the body through abrasions and other skin lesions. However, B. recurrentisis also capable of penetrating intact skin. As with R. prowazekii infections, body lice are killed as a result of infection with B. recurrentis. An intriguing history of human epidemics of louseborne relapsing fever is provided by Bryceson et al. (1970). Hippocrates described an epidemic of ” caucus,” or “ardent fever,” in Thasos, Greece, which can clearly be identified by its clinical symptoms as this malady. During 1727-1729, an outbreak in England killed all inhabitants of many villages. During the present century, an epidemic that spread from eastern Europe into Russia during 1919-1923 resulted in 13 million cases and 5 million deaths. Millions also were infected during an epidemic that swept across North Africa in the 1920s. Several major epidemics subsequently have occurred in Africa, with up to 100,000 fatalities being recorded for some of them. During and immediately after World War II, more than a million persons were infected in Europe alone. The only current epidemic of louse-borne relapsing fever is in Ethiopia, where 1000-5000 cases are reported annually, accounting for ca. 95% of the world’s recorded infections. Other smaller loci occur intermittently in other regions, such as Burundi, Rwanda, Sudan, Uganda, People’s Republic of China, the Balkans, Central America, and the Peruvian Andes. Resurgence of this disease under conditions of warfare or famine is an ominous possibility. Additional information on louseborne relapsing fever is provided by Bryceson et al. (1970).

TRENCH FEVER

Also known as five-day fever and wolhynia, trench fever is caused by infection with the bacterium Bartonella (formerly Rochalimaea) quintana. Like the two preceding diseases, the agent is transmitted by the human body louse. Human infections range from asymptomatic through mild to severe, although fatal cases are rare. Clinical symptoms are nonspecific and include headache, muscle aches, fever, and nausea. The disease can be cyclic, with several relapses often occurring. Previously infected persons often maintain a cryptic infection which can cause relapses years later, with the potential for spread to other persons if they are infested with body lice. Effective antibiotic treatment of patients involves administering drugs such as doxycycline or tetracycline. Lice become infected with B. quintana after feeding on the blood of an infected person. The pathogen multiplies in the lumen of the louse midgut and in the cuticular margins of the midgut epithelial cells. Viable rickettsiae are voided in louse feces, and transmission to humans occurs by the posterior-station route when louse bites are scratched. B. quintana can remain infective in dried louse feces for several months, contributing to aerosol transmission as an alternative route of transmission. Transovarial transmission does not occur in the louse vector. Infection is not detrimental to lice and does not affect their longevity. Trench fever was first recognized as a clinical entity in 1916 as an infection of European troops engaging in trench warfare during World War I. At that time, more than 200,000 cases were recorded in British troops alone. Between the two world wars, trench fever declined in importance but re-emerged in epidemic proportions in troops stationed in Europe during World War II. Because of the presence of asymptomatic human infections, the current distribution of trench fever is difficult to determine. However, since World War II, infections have been recorded in several European and African nations, Japan, the People’s Republic of China, Mexico, Bolivia, and Canada. Until recently, B. quintana was considered to be transmitted solely by body lice. However, several homeless or immunocompromised people, including HIV-positive individuals, particularly in North America and Europe, have presented with opportunistic B. quintana infections. This is manifested not as trench fever but as vascular tissue lesions, liver pathology, chronically swollen lymph nodes, and inflammation of the lining of the heart. Because some of these patients were not infested by body lice, an alternate mode of pathogen transmission may have been involved.

Pathogens Transmitted by Lice

HUMAN LICE AS INTERMEDIATE HOSTS OF TAPEWORMS

Occasionally humans become infested with the doublepored tapeworm (Dipylidium caninum). Although carnivores are the normal definitive hosts for this parasite, humans can be infested if they accidentally ingest dog biting lice (T. canis), which serve as intermediate hosts. Although this would appear to be an unlikely event, infants, especially babies playing on carpets or other areas frequented by a family dog, may touch an infested louse with sticky fingers which may then be put into their mouth, thus initiating an infestation.

VETERINARY IMPORTANCE

Several chewing lice and sucking lice parasitize domestic animals (Tables I, II). Although louse populations are usually low on these hosts, lice can sometimes multiply to extremely high numbers, particularly on very young, old, or sick animals. Often this is because hosts are unable to effectively groom themselves or they are immunocompromised. Except for the possibility of pathogen transmission, small numbers of lice typically cause little harm to the host. However, large numbers of lice can be debilitating by causing anemia, dermatitis, allergic responses, hair or feather loss, and other disorders. Lice also induce intensive host grooming, which can lead to the formation of hair balls in the stomach, especially in cats and canes. A few pathogens are lmown to be transmitted to domestic animals by lice (Table III). The most important of these are the viral agent of swinepox and the bacterial agents of murine haemobartonellosis and murine eperythrozoonosis, all of which are widely distributed. In addition to those listed in Table III, several pathogens have been detected in various species of lice, but there is no current evidence that lice are vectors of these organisms.

LICE OF LIVESTOCK

Although louse populations of a few hundred individuals commonly occur on healthy livestock, sometimes these numbers can reach into the thousands or, rarely, to more than a million per animal. It is under thc latter conditions that detrimental effects to the host occur. These include restlessness, pruritus, anemia, low weight gain, low milk yield, dermatitis, hide or fleece damagc, skin crusting or scabbing, and lameness. Large louse populations on domestic stock typically develop on juvenile, senile, sick, nutritionally deprived, or immunocompromised hosts.

Sucking louse infestations of cattle, such as those caused by the shortnosed cattle louse (Haematopinus eurysternus), the cattle tail louse (H. quadripertusus), and the longnosed cattle louse ( Linognathus vituli) (Fig. 4.7H), can cause serious damage to the host. This can be manifested as frequent rubbing of infested areas, hair loss, scab formation, slow recovery from disease or trauma, and low weight gain. Younger animals are typically more severely affected than older cattle. Mixed infestations of both chewing and sucking lice on cattle, or of both lice and nematodes, can affect weight gains more severely than single infestations. In single or mixed infestations, weight gains are typically lower in stressed cattle and those on low-nutrition diets. Sometimes, cattle sucking lice cause severe anemia, abortions, or even death. Irritation can be caused by small numbers of lice in sensitive cattle and usually results in frequent rubbing and subsequent hide damage. This rubbing also damages livestock facilities. Severely infested cattle often have patches of bare skin and a greasy appearance which results from crushing lice and their feces during rubbing. Under laboratory or confined conditions, at least three pathogens can be transmitted by cattle sucking lice, i.e., the causative agents of bovine anaplasmosis, dermatomycosis (ringworm) (Table III), and, rarely, theileriosis. The importance of cattie lice in transmitting any of these pathogens in nature is unknown but presumed to be low. Lice of horses and other equids typically do not greatly debilitate their hosts except when they are present in large numbers. Pruritus, hair loss, and coat deterioration may occur in severely infested animals. Horses with severe louse infestations are nervous and irritable; they typically rub against objects, kicking and stamping. Hair can be rubbed from the neck, shoulders, flanks, and tail base, resulting in an unkempt appearance that may affect the value of the horse. No pathogens are known to be transmitted by equid lice. Hog lice can imbibe significant volumes of blood from hogs, especially piglets, which often have larger infestations than adult pigs. Hog-louse feeding sites often cause intense irritation, leading their hosts to rub vigorously against objects, which can result in hair loss and reddened or crusty skin lesions. Haematopinus suis is a vector of the virus that causes swinepox (Table III), a serious and potentially fatal disease characterized by large pockmark lesions, mainly on the belly of infected animals. Some studies have implicated this louse as a vector of Eperythrozoon suis and E. parvum, causative agents of swine eperythrozoonosis, and of African swine fevervirus. However, transmission of these pathogens by lice appears to be rare, if it occurs at all, in nature. All species of lice that parasitize sheep and goats (Tables I and II) can cause debilitation, even when present in relatively small numbers, because of the potential

FIGURE 4.15 Fleece damage (wool slippage) in a sheep, caused by severe infestation with Linognathus africanus (Anoplura). (Courtesy of John E. Lloyd)

FIGURE 4.15 Fleece damage (wool slippage) in a sheep, caused by severe infestation with Linognathus africanus (Anoplura). (Courtesy of John E. Lloyd)

damage which they can cause to fleece and wool (Fig. 4.15). Some sheep develop hypersensitivity to the sheep biting louse (Bovicola ovis) (Fig. 4.16). This louse causes most sheep fleece devaluation worldwide and is the major cause of cockle, an economically disfiguring condition of sheep fleece that is particularly prevalent in New Zealand. Any increase in skin lesions or body rubbing in response to lice generally devalues wool or mohair. Different breeds of sheep and goats exhibit contrasting levels of resistance or tolerance to infestation by lice.

LICE OF CATS AND DOGS

Louse infestations of cats and dogs are most noticeable on sick or senile hosts. Under these conditions, louse populations can increase dramatically. Severe infestations of any of the four species involved usually cause host restlessness, scratching, skin inflammation, a ruffled or matted coat, and hair loss. The dog biting louse ( T. canis) is an intermediate host of the double-pored tapeworm (D. caninum) (Table III). Lice become infected when they ingest viable D. caninum eggs from dried host feces. The tapeworm develops into a cysticercoid stage in the louse, where it remains quiescent unless the louse is ingested by a dog, usually during grooming. In the dog gut, the cysticercoid is liberated and metamorphoses into an adult tapeworm. The dog sucking louse (L. setosus) has been shown to harbor immatures of the filarial nematode Dipetalonema reconditum, which parasitizes dogs, but whether or not these lice are efficient vectors remains unknown.

FIGURE 4.16 Sheep-biting louse (Bovicola ovis), showing prothorax and head with mandibles characteristically grasping a host hair: scanning electron micrograph. (From Price and Graham, 1997)

FIGURE 4.16 Sheep-biting louse (Bovicola ovis), showing prothorax and head with mandibles characteristically grasping a host hair: scanning electron micrograph. (From Price and Graham, 1997)

LICE OF LABORATORY ANIMALS

Some lice that parasitize laboratory animals initiate serious health problems by causing pruritus, skin lesions, scab formation, anemia, and hair loss. Others are vectors of pathogens that can cause severe problems in animal colonies (Table III). The mouse louse (P. serrata) is a vector of the bacterium Eperythrozoon coccoides, which causes murine eperythrozoonosis, a potentially lethal infection of mice that occurs worldwide. Infection of this blood parasite in mice can either be inapparent or result in severe anemia. Transmission of this pathogen in louse-infested mouse colonies is usually rapid. The spined rat louse (P. spinulosa) is a vector of the bacterium Haemobartonella muris, which causes murine haemobartonellosis (Table III), another potentially fatal blood infection that can cause severe anemia in laboratory rats. Laboratory and wild guinea pigs are parasitized by two species of chewing lice, the slender guineapig louse Gliricola porcelli) and the ovalguineapig louse ( Gyropus ovalis). Small numbers of these lice cause no noticeable harm, whereas large populations can cause host unthriftihess, scratching (especially behind the ears), hair loss, and a ruffled coat. Large infestations of the rabbit louse ( Haemodipsus ventricosis) can cause severe itching and scratching, which results in the host rubbing against its cage, often resulting in hair loss. Young rabbits are more adversely affected than are adults and may experience retarded growth as a consequence of infestation by H. ventricosis. The rabbit louse is also a vector of the causative agent of tularemia among wild rabbit populations (Table III).

LICE OF POULTRY AND OTHER BIRDS

Although louse populations may be very large on domestic fowl, including domestic chickens, turkeys, guinea fowl, pea fowl, and pheasants, no pathogens are known to be transmitted by these lice. Large populations often occur on birds with damaged beaks whose grooming ability is significantly impaired. The chicken body louse (Menacanthus stramineus) (Fig. 4.14) often causes significant skin irritation and reddening through its persistent feeding. Occasionally the skin or soft quills bleed from their gnawing and scraping action, with the lice readily imbibing the resultant blood. The shaft louse (Menopon gallinae) also causes significant losses to the poultry industry, including deaths of young birds with heavy infestations. Large infestations of chicken body lice, shaft lice, and other poultry lice may be injurious to the host by causing feather loss, lameness, low weight gains, inferior laying capacity, or even death. The vast majority of chewing lice are parasites of wild or peridomestic birds. Several of these lice are suspected vectors of avian pathogens. Some chewing lice of aquatic birds, including geese and swans, are vectors offilarial nematodes (Table III). Pet parrots, parakeets, budgerigars, and other birds also are subject to infestation by chewing lice, which is usually noticed only by the associated host scratching and by ruffled or lost feathers. Large populations of these lice can debilitate their hosts. Ranch birds, such as ostriches, emus, and rheas, are prone to similar adverse effects caused by their associated chewing lice.

PREVENTION AND CONTROL

Several techniques have been used in attempts to rid humans and animals of lice and louse-borne diseases. Preventing physical contact between lousy persons or animals and the items they contact, as well as various chemical, hormonal, and biological control mechanisms, comprise the current arsenal of techniques. Chemicals used to kill lice are called pediculicides. Clothes of persons with body lice should be changed frequently, preferably daily, and washed in very hot, soapy water to kill lice and nits. Washing associated bed linen in this manner is also advisable. Infested people should also receive a concurrent whole-body treatment with a pediculicide. Overcrowded and unsanitary conditions should be avoided whenever possible during outbreaks of human body lice and louse-borne diseases because it is under these situations that both can thrive. Crab lice can often be avoided by refraining from multiple sexual partners and changing or laundering bed linen slept on by infested persons. Pediculicides should be applied to the pubic area and to any other infested body regions. To reduce the spread of head lice, the sharing of combs, hats, earphones, and blankets, especially by children, should be discouraged. Often, parents of children with head lice are notified to keep youngsters away from school or other gatherings until the infestation has been eliminated. If the parents are also infested, this can further involve ridding the entire family of lice to prevent reinfestations. Various pediculicidal shampoos, lotions, and gels are widely available for controlling head lice. These treatments typically kill all nymphal and adult lice, but only a small proportion of viable louse eggs. Therefore, treatments should be repeated at weekly intervals for 2-4 weeks in order to kill any recently hatched lice. Hatched or dead nits which remain glued to hair may be unsightly or embarrassing, and these can be removed with a fine-toothed louse comb. Louse combs have been used, in various forms, since antiquity to remove head lice (Mumcuoglu 1996). A wide range of pediculicides is commercially available. Although its use is now banned in many developed countries, the organochlorine DDT is widely used, especially in less developed countries, for controlling human and animal lice. Several alternative pediculicides, such as lindane, chlorpyrifos, diazinon, malathion, permethrin, or pyrethrins, are currently used throughout the world. Pediculicides can be used in powders, fogs, or sprays to treat furniture or premises for lice. Several general parasiticides show promise as pediculicides. Avermectins such as abamectin, doramectin, and ivermectin can kill human body lice and livestock lice. Prescribed doses of these compounds can be administered orally, by injection, or as topical applications of powders, dusts, and pour-ons. However, many of these compounds have not yet been approved for use on humans. The development of novel control agents for lice is a constant process because resistance to various pediculicides has developed in lice in many parts of the world (Burgess 1995, Mumcuoglu 1996).

Lice of livestock can be controlled by both husbandry practices and chemical intervention. Providing a high-energy diet, especially to cattle, can be an effective louse control strategy. If possible, it is important to keep animals in uncrowded conditions and to spot-treat or quarantine any infested individuals until they have been successfully deloused. Various formulations and applications of pediculicides are typically used to control lice on livestock. Insecticidal dusts, powders, sprays, dips, ear tags, tail tags, resin strips, gut boluses, collars, pour-ons, lotions, and injections are widely used products. Infested animals should be treated twice weekly for 2-4 weeks. Insecticidal dust bags or back rubbers can be used as selfdosing rubbing stations for cattle and other livestock. Because louse populations on livestock are typically greater during the winter months, pediculicides are usually best applied to them in the late fall. Fall systemic treatments of cattle for both lice and bots are often administered. Shearing wool from sheep removes up to 80% of the lice present on infested animals. Pets, laboratory animals, and poultry can be treated for lice in several ways. Pets such as dogs and cats can be dipped or bathed with a pediculicidal lotion or shampoo. Various oral or topically applied insecticides used for controlling fleas on pets also are efficacious against lice. Similarly, flea combs also remove lice from pets. Poultry and laboratory animals can be treated with pediculicidal dusts or sprays. Although host treatment is most efficacious, bedding materials and cages can also be treated. Insecticidal feed additives are also available. Insecticideimpregnated resin strips can be added to cages of poultry or laboratory animals to control lice. The bacterium Bacillus thuringiensis and the nematodes Steinernema carpocapsae and S. glaseri, which are effective biological control agents against numerous arthropods, can also be used to kill livestock lice. Some juvenile hormone analogs and insect growth regulators such as diflubenzuron have similarly shown promise as pediculicides. With respect to louse-borne diseases, vaccines have been developed only against epidemic typhus, and none is completely safe or currently approved for widespread use. The live attenuated E-strain vaccine has been administered to humans, particularly in certain African nations, in attempts to quell epidemic typhus outbreaks. However, this vaccine actually caused disease in some patients and did not always prevent subsequent infection.

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