Phthiraptera: India
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Phthiraptera
Introduction
The Lice (the 'chewing-, bird-, biting-, or feather-lice', and the 'sucking-lice' are dorso-ventrally flattened obligatory ecto-parasitic insects on the warm blooded vertebrates, viz., birds and mammals. While the chewing-lice are parasitic on both birds and mammals, the sucking-lice are exclusive to mammals. There are about 2900 (Clay, 1974) to 3000 species (pilgrim, 1970) of chewing-lice, and about 450 species of sucking-lice.
The lice live in the plumage of the birds, or pelage of mammals, and well adapted to live in the micro-environment of the feather, or hair cover, which not only offer them shelter, but also food, and the parasites are thus exposed very little to the external or macroclimatic fluctuations to which the hosts are exposed. They live in one zone, feed in another, and oviposit in yet another, depending on the species involved (Ash, 1960). Their food chiefly consists of the' feathers, hair sebum, serum and blood in case of the chewing-lice, and serum and blood in case of the sucking-lice. As the popular name• indicates those species wi~ typical mandibulate or masticating type of mouth parts are called 'chewing-lice', and those species adapted for typical piercing and sucking type of mouth parts are included under 'sucking-lice' However, this is not exclusively true from a functional angle, since some chewing-lice with well developed mandibles like the Ricinidae developed a modified piercing and sucking apparatus, and the Haematomyzidae have mandibles. rotated 1800 so that their cutting or dentated edges face away from one another and probably at the most useful for anchoring to the host body, and the species live more by sucking.
The sexes are separate, more often females oUblumber males, and in quite a few instances the males are even unknown, and probably the species are parthenogenic. There are occasional predominance of males over females, and at least in one case of Piagetiella parasitic on the grey pelican, it was attributed to the adverse conditions faced by the host and its health was affected (Madhav, Ramana Rao, &Lakshminarayana, 1990). The males also sometimes show extreme sexual dimorphism. Not only asymmetry played a dominant part in the sexual isolation amongst the sympatric species, there are certain genera of chewing-lice where only exclusively asymmetrical forms are known (Lakshminaryana, 1977, 1979 &Lakshminarayana &Emerson, 1971, 1978). The life-cycle is simple with an e~g, three nymphal instars , and the adult.
The lice cause dermatitis, spread diseases like rickettsiasis, murine typhus, etc., causing annoyance to the host during the course of feeding or movement, known to effect the egg production, and in case of heavy infestations the hosts not only present sickly appearance, but also become exposed to secondary infections. Those species feeding on the hair may damage the coat, and those that feed on the serum and blood are capable of producing toxaemia, inject toxins and other germs like viruses and bacteria. Some of the chewing-lice act as intermediate hosts, for example, the swift filarial worm Filaria cypseli is transmitted by Dennyus minor, dog cestode by Trichodectes canis, transmission of typhus in guinea pig by Trimenopon hispidum, and infectious anaemia of horses by Werneckiella equi. Emerson (1973) states that the eastern equine enceph lomyelitis virus and Bedsonia organisms in pheasants were also found in the chewing-lice of the respective hosts.
The role of the chewing-lice in disease transmission is however, little explored, and with the increased interest in the wild-life conservation and epizoology and diseases, we should have better understanding on the role of lice in disease transmission.
They have no intermediate hosts unlike many other parasites, and the transmission from one host to the other is possible only between the sexes during feeding, roosting, or mating, between the offspring and the parent; also possible between the prey and the predator, or foster parent and the brood parasite, during dust baths, and rarely by phoresy. However, it is seen that the transmission is direct, and the non-establishment of species on unrelated hosts, prey and predator (majority cases), the foster parent and its brood parasites indicate their host specificity. In fact, the lice and their hosts evolved hand in-hand, and therefore, the study of evolution in one reflects that of the other. This has been taken as advantage by the parasitologists and they suggested host relationships and host distribution.
Hopkins (1949, 1957) reported that the ancestors of Amblycerophtihrina (the most -primitive member of the chewing-lice) possibly began their life as ecto-parasites of vertebrates during the Triassiac Period (225-190 million years ago), and parasitized the early birds and mammals, and possibly their reptilian ancestors. True Amblycerophthirina possibly evolved at least in Jurassic (190-135 m.y. ago), Ischnocerophthirina type' of Chewing-lice might have been in-existence since early Cretaceous (135-65 m.y. ago), or even in late Jurassic, and the Siphunculophthirina (sucking-lice) not later than middle Cretaceous. The evidence of Rhynchopthirina chewing-lice is too scanty, but it must have evolved earlier than Eocene Epoch (54-38 m.y. ago), and possibly existed even in Cretaceous Period.
Classification
A lot of confusion prevailed regarding the classification, priorities and authorship of various suprageneric taxa in the lice as a whole. A good review on the nomenclatural problems, priorities, and authorships was presented in Lakshminarayana (1970, 1976). Majority of phthirapterologists• now accept Phthiraptera the name of the Order. The former suborders Amblycera, Ischnocera, Rhynchophthirina of Mallophaga (chewing-lice), and Siphunculata (Anoptura, sucking-lice) emended as Amblycerophthirina, Ischnocerophthirina, and Siphunculophthirina on par with Rhynchophthirina Ferris, which also indicate their natural relationship (Lakshminarayana, 1970, 1976) are now considered as suborders of Phthiraptera. Hopkins &Clay (1952) included the families Menoponidae (incI. Trinotonidae), Laemobothriidae, Ricinidae, Boopiidae, Trimenoponidae, and Gyropidae under Amblycerophthirina, Philopteridae, Trichodectidae, and Heptapsogasteridae under Ischnocerophthirina and the monotypic family Haematomyzidae under Rhynchophthirina. Mukerji &Sen-Sarma (1955) suggested to separate -the suborder Rhynchophthirina as RhYnchophthiraptera. Carriker (1960) proposed a new family Trochiliphagidae with type genus Trochiloecetes. Clay (1962) changed it as Trochiloecetidae. Keler (1964) revised earlier classification. Emerson &Price (1976) erected the family' Abrocomophagidae under Amblycerophthirina. Lyal (1985) thoroughly revised the family Trichodectidae.
Ferris (1951) revised his earlier series of monographs on sucking-lice (1919-1933) and divided them under six families viz., Echinophthiriidae, Haematopinidae, Pediculidae, Hoplopleuridae, Neolinognathidae, and Linognathidae. Hopkins (1949, 1957) gave a thorough account of lice of mammals and the host relationships. Johnson (1964) removed the subfamily Pedicfninae from Hoplopleurid~e; he also proposed (1969) a new family Haemophthiridae to accommodate the solitary species Haemophthirus galeopitheci"removing it from Hoplopleuridae. Kim (1970) proposed a seven family classification transferring Pedicininae back to Hoplopleuridae.
The present classi'fication is as follows: Order Phthiraptera; Suborder Amblycerophthirina (Boopiidae, Menoponidae s.l., Trinotonidae, Laemobothriidae, Ricinidae, Trochiliphagidae (frochiloecetidae), Gyropidae, Abrocomophagidae, Trimenoponidae), Suborder Ischnocerophthirina (Philopteridae, s.l., Heptapsogasteridae, and'Trichodectidae) and the Suborder Rhynchophthirina (Haematomyzidae) under chewing-lice. The Suborder Siphunculophthirina (Siphunculata or Anopluta auct.) (Echinophthiridae, Haematopinidae, Linognathidae, Pediculidae, Neolinognathidae, Hoplopleuridae) (see fig. 1).
Historical Resume
i) Pre-1900
Denny (1842) gave a historical account of lice from the Biblical times. There are other three works on the pre-Christian Era viz., Herakleitos Ephesius (500 B.C.), Aristotle (350 B.C.), and Diophanes of Bithynia (100 B.C.).
Lice were however, been known from much earlier times in India. In Manava Dharma Sastra (Vedic Period), flies mosquitoes, lice and bugs were classed amongst the animals that bred in sweat (svedaja). Umasvati's classification in Tatvarthadigama (circa 40 A.D.) in which the lice were classed Trapusarja and Karpasathika on the basis of their sense organs. In the Sangam literature of South India (a period assigned between 3000 B.C. to 1915 A.D., according to some and from 4 A.D. to 8 A.D. according to others), are found plentiful reference to various insects like ants, bees, wasps, dragonflies., white-ants, and lice as well as scorpions, besides the habits and habitats of other animals like birds. Emperor Asoka (273-232 B.C.) established a number of veterinary hospitals whose inmates include various birds and mammals and the doctors in-charge of these hospitals were said to have a good knowledge of their scourges. Moghul Emperors seem to have a fair knowledge of birds. Abul Fazl, a courtier to Akbar, is said to have referred in his works, that Kashmir vallery was infested with such undesirables like gnats, -flies, fleas, and lice, etc.
In Ancient India, several birds like domestic hen, geese, pigeons and doves (for carrying post), peafowl 'parrots' (probably paraquets), and mynahs, and various mammals like cattle, elephants, horses, etc, were domesticated, their habits were well known, diseases were carefully observed, and treatises like Gajasastra, Aswasastra, samhitas like Charaka and Susruta were wri.tten. These treatises included various diseases their causative organisms including lice, and their cures. Lice even entered in proverbial lIterature. For example, there is a proverb in Telugu that !"Penuku (lice) pettanamichchina (if power is given) tala tega korukun (bites the head completely)"!, thereby meaning, if power is given to undesirables, they misuse it.
The fust scientific report of lice on Indian birds apparently dates back to J. C. Fabricius (1775) who described the habitat of Pediculus vulturis (Laemobothrion vult uris as Indiae Orientalis vulturibis. It was followed by Pediculus tantali (Ardeicola tantali) from the painted stork Tantalus leucocephalus (Ibis leucocephalus (pennant» from Tranquebar (Tamil Nadu) in 1798 described Lipeurus himalayensis (now Reticulipeurus himalayensis) from the western horned pheasant, Tragopon hastingii (T. melanocephalus). Few other references to the Indian species of chewing-lice are found in Giebel (1874), Piaget (1869) Walker (1871) and Richtar (1871).
ii) 1901-1947
Kellogg (1908), Gaiger (1910, 1915) Piaget (1915) and Paipe (1912) were based from material collected from animals in the European zoologi.cal gardens, or museum skins. Kellogg &Paine (1914) were the first to study the material collected actually from India by' the Indian Museum, Calcutta (now in ZSI). This was followed by another short list by Kellogg &Nakayama (1915). Notable contributions made before 1947 are as follows, chiefly by westerners: Ansari (1943-1944), Bhattacharjee (1939), Clay (1936, 1938, 1940, 1947), Clay &Meintertzhagen (1937, 1938 a-b, 1941), Conci (1941, 1942), Gaiger (1910, 1915), Overgaard (1943), Quadri (1935, 1936 a-c), Salim Ali (1927, 1936), Sen (1942), Thompson (1937, 1938, 1939, 1940 a-c), and Waterston (1928).
iii) 1948-1990
Agarwal (1967), Agarwal &Gupta (1970), Agarwal &Saxena (1977, 1978 a-b, 1979 a-b, 1980 a-b), Ansari (1947, 1951 a-b, 1954, 1955 a-g, 1956 a-i, 1957 a-e, 1958,1959, 1967), Arora &Chopra (1957, 1959), Brelih (1965), Carriker (1976), Chopra (1969), Clay (1947, 1949 a, 1958 a-b, 1959, 1962a, 1963, 1965a, 1966 a-b, 1970b, 1973, 1974a), Clay &Hopkins (1950, 1951, 1954, 1960), Dalgleish (1969), Dhanda (1961), Elbel (1967), Elbel &Price (1970, 1973), Emerson (1954), 1955, 1965, 1971, 1972 a-b, 1973), Emerson &Elbel (1956, 1957 a-c), 1961, Emerson &Price (1968, 1974), Guimaraes (1974), Hajela (1970), Hajela &Tandan (1967, 1970), Hopkins &Clay (1952, 1953, 1955), Hopkins &Timmermann (1954), Keler (1939, 1958, 1960, 1964, 1971), Klockenhoff (1969 a-b, 1973, 1980, 181), Klockenhoff &Schirmers (1976)
Klockenhoff, Rheinwald, &Wink (1973), Kumar &Tandan (1968, 1971), Lakshminarayana (1968 a-b, 1969, 1970 a-d, 1972 a-b, 1973 a-b, 1975 a-b, 1976, 1977 a-c, 1979 a-c, 1980, 1981, 1982 a-b, 1983, 1984, 1985, 1986, 1987, 1988, 1990), Lakshminarayana, Vijayalakshmi &Talukdar (1980), Ledger (1970, 1971), Lyal (1985), Madhav, Ramana Rao &Lakshminarayana (1986 a-b, 1990), Mukerji &Sarma (1955), Nelson &Price (1965), Price (1964 a-b, 1965, 1967, 1968, 1970 a, 1970 b-c, 1971, 1974, 1975, 1977), Price &Beer (1963 a-d, 1964, 1965 a-c), Price &Elbel (1969), Price &Emerson (1966,a,b 1967, 1974, 1975, 1977), Price &Beer (1963 a-d, 1964, 1965 a-c), Price &Elbel (1969), Price &Emerson (1966a, b, 1967, 1974, 1975, 1977), Rai (1977, 1978), Rai &Lakshminarayana (1980), Rakshapal (1959), Rao, Khuddus, &Channabasavanna (1975), Rheinwald (1968), Ryan &Price (1669), Saxena &Agarwal (1977 t 1978, 1979, 1980 a-d), Scharf &Price (1977), Sen &Fletcher (1962), Seneviratna (1963), Somadder &Tandan (1970), Srivastava (1977, 1978), Tandan (1951, 1952 a-b, 1955, 1958, 1963, 1978, 1973, 1976), Tandan &Clay (1971), Tandan &Kumar (1969), Tendeiro (1958, 1962, 1963, 1965 a-c, 1667 a, 1969 a-b, 1972, 1973 a, 1974), Thompson (1948 a-c, 1950 a-c, 1948), Timmermann (1952, 1954 a-c, 1955, 1962, 1965), Ward (1955), and Werneck (1950 have contributed on the subject.
The full references to the above works may be found in Keler (1960, Mitt. zooI. Mus. Berl.,. 36 (2) : 146-403), Eichler &Zotorzycha (1969, Angew, Parasit., 10 (1) : 53~60), Eichler, Ribbeck &Zotorzycka (1973, Mitt. zool. Mus. Berl., 49 : 423-461), and Lakshminarayana (1972, Angew. Parasit., 13 (3) 169-178; 1973, Angew. Parasit., 16(1) : 28-30; 1986, Misc. publ. Oee. pap. Rec. zool. Surv. India, 81 : 1-63).
Biological Studies
The distribution of parasites (specially obligatory ones like lice) depends mostly on the geographical distribution of the host, and synchronises with the latter. Thus, studies on the basis of hosts is more relevant and scientific. Most of the Eueropean and American workers based their studies on the collections made during the colonial rule,• specially by Col. Dr. Meinertzhagen, collections made by the SEA TO in the south-east Asia, few collections made by individual scientific groups either from India and adjacent countries or from their zoo hosts. Dr Ansari worked in this region who based his studies on collections made from Delhi and Izzatnagar (U.P.) in India and Pakistan and Bangladesh. Agarwal &his coworkers, Arora &Chopra, Tandan, Dhanda and coworkers and Qadri from U.P. and Punjab; Lakshminarayana, Rai, Vijayalakshmi and Talukdar from Z.S.I. from Eastern India, Nepal, Andaman &Nicobar, Lakshminarayana, Madhav &Ramana Rao from Andhra Pradesh, worked on lice fauna.
Estimation of Taxa
A perview of the avian lice indicates that a greater number of genera of chewing-lice are parasitic on aquatic &semi-aquatic birds than terrestrial ones, as follows: No.of Avian fauna of the World Host Genera Parasite Genera Aquatic,semi -aquatic,shore birds &waders 275 71 25.8% Terrestrial birds 1786 170 9.5% Schematic representation of the suborders and families of Phthiraptera Lakshminarayana (1986) presented the country-wise distribution of the chewing-lice as follows: Country Genera Species
Lakshminarayana (1986) also stated that the reported species from India, Afghanistan, Bangladesh, Bhutan, China, (adjoining areas to India), Myanmar (Burma), Nepal, Pakistan and Sri Lanka are as follows: Families No. of Genera Species
Our knowledge on Indian mammalian chewing-lice is practically nil, confined only to about 9 host species. Hopkins (1949) while reviewing the lice on. mamm~s stated that we know considerable number of lice from Ethiopian Region, but our knowledge of lice from Oriental Region is practically nil, although, the Oriental Region is equally rich in mammals. This statement holds true even now. Unless greater interest on wild life epizoology is generated and diseases of the wild life are taken up simultaneously with its conservation programme, the position is not likely to change. Classified Treatment Systematics
Keys for the identification of the suborders and families from India are given in Lakshminarayana (1970,1986). Keys for the identification of genera and species are also given in Blagoveshthenskii (1964,1967), Clay (1947,1976,1977,1978). The valid genera and species alongwith type-hosts and fust references can be found in Hopkins (1949,1957), Hopkins &Clay (1952,1953,1955), Lyal (1985), Price (1973), Price &Emerson (1966,1967) and the list of species from India and adjacent countries in Lakshminarayana (1979,1982).
Morphology
Mukerji &Sen Sarma (1955) studied the morphology of the elephant louse, H aematomyzus elephantis in detail. Rao et al. (1975) studied the mouth parts of the chewing-Jice. A glossary of taxonomic characters (Lakshminarayana, 1985) were given. The role of asymmetry was discussed in Lakshminarayana (1973a,b, 1977a,1979b), and Lakshminarayana &Emerson(1971,1978). The trends in the evolution of the sitophore sclerite and" male genitalia were discussed in the Laemobothrion-complex (Lakshminarayana, 1970b).
Anatomy
Mukerji &Sen Sarma (1955) studied the anatomy of the elephant louse, Haematomyzus elephantis. Srivastava (1974) studied the poultry louse, Numidilipeurus lawrensis tropicalis which was also studied in detail by Agarwal &Saxena, (1978, 1979b, 1980). Saxena &Agarwal, (1980a¬c) dealt with the crop, teeth, tracheal, vascular and genital systems.
Histology
The histology of various organs in Haematomyzus elephantis was discussed in Mukerji &Sen Sarma (1955). Srivastava (1974), and Saxena and Agarwal (1979,1980d) also discussed on this problem with reference to N.J. tropicalis. Agarwal &Saxena(1977) worked out the histology of peri-oesophageal nephrocytes in some lice.
Behaviour
Rakshapal (1959) observed the behaviour of the pigeon louse, Columbicola columbae. Agarwal &Saxena (1980) studied the feeding behaviour of N.J. tropicalis. Clay (1949c) discussed the behaviour and its impact on the morphological evolution. Lakshminarayana &Emerson (1971,1978) discussed on the probable changes brought about in two sympatric species of Goniodes consequent to the evolution, isolation, and reunion of two species of its host genus viz., Pavo in the Oriental Region. Lakshminarayana (1973b) also attributed the morphological changes due to feeding habit in the head asymmetry in Struthiolipeurus. It was contended that an ancestral form originally parasitic probably on Falconiformes, secondarily parasitized Struthioniformes and Rheiiformes, developed the asymmetry due to the feather structure in the latter group of hosts.
Bionomics
Ansari (1944a,b) studied the bionomics of the lice of domestic chicken. Arora &Chopra(1957,1959) studied the biology of NJ.tropicalis. Agarwal(1967),studied the bionomics of FalcoJipeurus Irater. Agarwal &Saxena(1978,1979a) studied the seasonal dynamics of N i.tropicalis and its feeding habits (Agarwal &Saxena, 1980a). Rai &Lakshminarayana (1980) discussed techniques for in vitro studies on the chewing-lice. Evaluation of secondary infestations on the basis of immature stages was discussed in Lakshminarayana (1972a). Madhav et al(1990) studied sex ratio and importance of Piagetiella on grey pelican-population control.
Development
Ansari (1954) studied the pre-imaginal instars of the chewing-lice and applied the growth principles. Agarwal(1967) studied the development of Falcolipeurus Irater. Agarwal &Gupta (1970) studied the effect of low temperatures on the viability of the eggs and development in F. frater. Rai &Lakshminarayana (Op.CiL) gave method ofrearing artificially. Cytotaxonomy Practically no work is available on this aspect. However, Manna (1990) reported the chromosomal numbers of six species.
Expertise India
Z.Sl.
K. V. Lakshminarayana, Zoological Survey of India, Southern Regional Station, 10Q, Santhome High Road, Madras 600 028. [Taxonomy, Host Parasite relationship, Evolution.]
Elsewhere
G. P. Agarwal, Department of Zoology (parasitology), Benaras Hindu University, Varanasi 221 005.[Bionomics, Histo-morphology] A. K. Saxena, Depu. of Zoology, Pt. L.M.S. College, Rishikesh 249 201, Debra Dun(U.P.) [Bionomics, Risto-morphology] B. K. Tandon, Department of Zoology, Lucknow University, Lucknow 226 001 (U.P.) [Taxonomy, Host Parasite relationship]
Abroad
Dale H. Clayton, Committee on Evolutionary Biology, University of Chicago, Chicago, Illinois 60637.(U.S.A) [Taxonomy &Evolution.] Eberhard Mey, Naturhistorisches Museum der Staatlichen Museum, Heidechsburg Rudolstadt PF48/51Rudolstadt/Thur 6820(Germany ) [Taxonomy] Wd. Eichler, Museum fUr Naturkunde an der Humboldt Universitat, 104 Invalidenstrasse, Berlin (Germany) [Taxonomy, Bionomics, Evolution Host Parasite relationship Control] K. C. Emerson, 560 Boulder Drive, Sanibel, FL 33957 (U.S.A) [Taxonomy, Evolution, Host Parasite relationship] J. A. Ledger, Department of Entomology, South African Institute for Medical Research, Johannesburg, (S.Africa) [Taxonomy, Evolution] H. W. Ludwig, Professor am Zoologischen, Institut der Universitat Heidelberg, Heidelberg (Germany) [Taxonomy, Control] R. L. C. Pilgrim, Zoology Department, University of Canterbury, Private Bay, Christ Church (New Zealand) [Taxonomy] Robert Timm, Museum of Natural History &Department of Systematics &Ecology, University of Kansas, Lawrence, Kansas 66045 (U.S.A) [Taxonomy] Roger D. Price, Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, (U.S.A) [Taxonomy,Evolution, Host Parasite Relationship] Ronald A. Hellcnthal, Department of Biological Sciences of Notre Dame, Notre Dame, In 46 556. [Taxonomy] Jadwiga Ztotonycka, Department of Parasitology, University ofWroclaw (poland) [Taxonomy, Evolution, Host Parasite Relationship, Control] Richardo L. Palma, National Museum of New Zealand, P.O. Box 467, Wellington, (New Zealand)[Taxonomy]. '
Selected References
Clay, T. 1951. An Introduction to a classification of the avian Ischnocera (part I). Trans. R. ent. Soc. Lond. 102: 171-194. Clay. T. 1969. The Amblycera (phthiraptera:Insecta). Bull. Br. Mus. nat. Hist. (Ent.) 25 (3): 73¬98, 9figs, 5 pIs.
Eichler, Wd. 1963. "Phthiraptera 1. Mallophaga" In Bronn's Klassen und Ordnungen des Tierreichs.-/Akad. Verlagegesellschaft Geest &Portig, K.G./,Leipzig, 5 (3):7b.vii¬29Opp., 150 figs. Hopkins, G.R.E. &Clay, T. 1952. A Checklist of Genera and Species of Mallophaga. Br. Mus. nat. Hist. London : 362 pp.
Hopkins, G. H. E. &Clay, T. 1953,1955. Additions and corrections to the Check-list of Mallophaga. Ann. Mag. nat. Hisl., (12) 6 : 434-448; II. Ann. Mag. nat. Hist., (12) 8 : 77-80.
Keler, S. 1960. Bibliographie der Mallophagen. Mitt. zool. Mus. Bert., 36 (2) : 146-403. Lakshminarayana, K. V. 1979-1982. A Synoptic list of Mallophaga sens.lat. (Phthiraptera:Insecta) from India and adjacent countries together with host &Regional indices. Rec. zool. Surv. India, 75 : 39-201; 80 : 61-83. Lakshminarayana, K. V. 1985. Glossary of Taxonomic characters for the study of chewing-lice. Misc. Publ. Occ. pap;Rec. zool. surv. India, 82 : 59 pp.
Lakshminarayana, K. V. 1986. Data book for the study of the Chewing-lice (phthiraptera:lnsecta) in India and Adjacent countries Misc. Publ. Occ. pap; Rec. Zool. Surv. India, 81:63 pp. Lyal, C. H. C. 1985. A cladistic analysis and classification of tricholodectid mammal lice (phthiraptera: Ischnocera). Bull. Br. Mus. nat. Hist. (Ent.), 51 (3) : 187-346.
