Lice and Fleas ( Phthiraptera and Siphonaptera ) Chapter 13 . 4

Abstract A total of 31 Phthiraptera species alien to Europe are listed. Th ey include 24 chewing lice and seven sucking lice of 12 diff erent families. Th e families Goniodidae (Ischnocera) and Menoponidae (Amblycera) largely dominate the alien entomofauna of chewing lice. Asia is the major supplier of alien Phthiraptera which are mostly associated with poultry farming, game birds, guinea pigs and invasive alien mammals. Th e recent period did not show any acceleration in alien arrival in Europe. Alien fl eas include six species in the families Pulicidae and Ceratophyllidae. Th ree of them are primarily associated with rats and are capable of transmiting major human diseases such as the bubonic plague and the murine typhus.


Introduction
Phthiraptera (lice) and Siphonaptera (fl eas) are obligate ectoparasitic insects of birds and mammals, including humans.Some are of high importance for human and animal health because they cause itches and skin infection, and transmit serious diseases, e.g. the head louse (Pediculus capitis De Geer), the crab louse (Phtirus pubis (L.)), the cat fl ea (Ctenocephalides felis felis (Bouché)), the rat fl ea (Xenopsylla cheopis (Rothschild)) or the human fl ea (Pulex irritans L.).Although many of these are of unknown origin, they are probably allochtonous in Europe, having arrived in ancient times with their hosts (Mey, 1988;Beaucournu and Launay, 1990).Th us, Pulex irritans was shown to have been present in Europe since the Bronze Age at least, having been found in remains of lake dwellings in the French Jura, dating back to 3100 B.C. (Yvinec et al 2000).
Only the species considered as possibly neozoans*, i.e. 27 lice and six fl eas, were originally included in the DAISIE database.Four further species have subsequently been added to the list of alien species and this review is therefore based on 31 species.
Although a large part of these alien species were recorded in Europe for the fi rst time at the end of the 19 th century, many probably came much earlier; the exact date of arrival remaining unclear in nearly all cases.

Phthiraptera
Lice are exopterygotes* of birds and mammals.Most species are host-specifi c but others are rather polyphagous.Th ey spend their entire life on their host animal, feeding on epidermal tissue debris, parts of feathers, blood or sebaceous secretions.Until recently, they were divided into two orders, Anoplura (sucking lice) and Mallophaga (chewing lice), but they are presently grouped into a single order, Phthiraptera (Barker et al 2003;Price et al 2003).Th e order Phthiraptera comprises about 5,000 described species present in four sub-orders, Anoplura (543 spp. on mammals), Amblycera (ca.1360 spp. on birds, mammals and marsupials), Ischnocera (ca.3080 spp. on birds and mammals) and Rhynchophthirina (3 spp. on elephants and warthogs), this latter group being not present in Europe (Smith 2003).
A total of 31 Phthiraptera species alien to Europe have been listed here, including 16 species known to be of exotic origin and 14 cryptogenic species, to be compared to the 691 species considered as native to Europe included in Fauna Europaea (Mey 2005).Th ey include 24 chewing lice belonging to 8 diff erent families and 7 sucking lice belonging to 4 diff erent families (Table 13.4.1).Th ree of the families have no representatives in Europe (Gliricolidae, Gyropidae, Trimenoponidae; all in the Amblycera suborder).Th e families Goniodidae (Ischnocera) and Menoponidae largely dominate the alien entomofauna (Figure 13.4.1).In a number of families, the arrival of aliens has largely modifi ed the composition of the total entomofauna currently present in Europe.
In contrast to the trends reported in other arthropod groups, the majority of the alien lice were fi rst observed in Europe during the 18 th and 19 th century (18 species out of 31-58.1%),although they probably arrived much earlier with their animal host, in most cases a domestic species.Th e recent period did not show any acceleration in alien arrival in Europe with only 4 species (12.9 % of the total species) newly observed during the period 1975-2007.Eight out the 17 alien species of known exotic origin came from Asia (47.0 %), with earlier arrival dates than those from North America (4 spp.; 23.5 %) or South America (4 spp.).
Several chewing lice of cryptogenic origin are important pests of poultry farming, in particular Menopon gallinae, Goniocotes gallinae and Eomenacanthus stramineus (Sychra et al 2008).Other species parasitize pheasants (Phasianus spp.) and came with their host from Asia, such as Goniocotes chrysocephalus, Lagopoecus colchicus, Lipeurus maculosus, Uchida phasiani, Zlotorzyckella colchici (Kopocinski et al 1998).Chewing lice parasitising mammals in Europe are listed in Mey (1988).Some species are known to be of alien origin, such as the three South American species, Gyropus ovalis, Gliricola porcelli and Trimenopon hispidum, arriving in Europe with guinea pigs (Cavia porcellus L.) and causing scratching, loss of hair, and scabs to domestic and laboratory animals.Other species worth mentioning are the cryptogenic dog louse, Trichodectes canis, and the sheep louse, Bovicola ovis, which cause pruritus and skin infections such as eczema to their host animal.Finally, a few species are associated with invasive alien mammals, such as the South American Pitrufquenia coypus on coypu (Myocastor coypus (Molina)); (Laurie 1946;Newson and Holmes 1968) and the North American Trichodectes (Stachiella) octomaculatus on raccoon (Procyon lotor (L.)); (Hellenthal et al 2004).
In addition, Haemodipsus ventricosus (Denny) which lives on rabbits (Oryctolagus cuniculus L.) can be considered as alien in Europe, originating, as its host, from the Iberic pensinsula (Durden and Musser 1994).

Siphonaptera
Fleas are holometabolous insects whose adults must feed on blood of mammals and birds in order to reproduce.Larvae feed on organic matter, often in the host's nest.In the DAISIE database, six fl eas are listed as alien to Europe, including 5 species known to be of exotic origin and 1 cryptogenic species, in comparison to the 260 species considered as native to Europe (Soledad Gomez Lopez 2005) (Table 13.4.1).Th e aliens belong equally to two families, Pulicidae and Ceratophyllidae, whereas the latter family largely dominates the native entomofauna.Th ree of these fl eas have rats as their main host (Beaucornu and Launay, 1990).Th e tropical rat fl ea, Xenopsylla cheopis, probably originates from the Nile area (Beaucornu 1999).It became synanthropic in most of Southern Europe where it could not survive before because of large temperature variations between summer and winter within human habitats (Beaucornu 1999).X. brasiliensis, originates from tropical Africa and invaded the Canary islands (Beaucornu and Launay, 1990); it has also been found sporadically in port areas and elsewhere, e.g. it was recorded from Wales in the 1950s (Hopkins and Rotschild 1953).Th e third species, Nosopsyllus fasciatus, is a temperate species from Asia.Rat fl eas are also able to feed on other mammals, including humans, to which they can transmit the bubonic plague by carrying the bacteria Yersinia pestis (Audouin-Rouzeau, 2003).Xenopsylla cheopis is also a vector of another human disease, the murine typhus fever caused by the bacteria Rickettsia typhi (Beaucournu and Launay, 1990).Th e North American species Orchopeas howardi is found on the grey squirrel (Sciurus carolinensis), an invasive rodent in Europe (Keymer, 1983).

Figure 13
Figure 13.4.1.Relative importance of the Phthiraptera and Siphonaptera suborders and families in the alien and native fauna in Europe.Families of Phthiraptera are presented per suborder in a decreasing order based on the number of alien species.Species alien to Europe include cryptogenic species.Th e number over each bar indicates the number of species observed per family.

Table 13 .4.1.
List and characteristics of the Phthiraptera and Siphonatera species alien to Europe.Status: A ,