Apeer-reviewed open-access journa BioRisk 4(2): 833-849 (2010) i d op ji 1] doi: 10.3897/biorisk.4.65 RESEARCH ARTICLE B | O R IS k www.pensoftonline.net/biorisk Lice and Fleas (Phthiraptera and Siphonaptera) Chapter 13.4 Marc Kenis', Alain Roques? | CABI Europe-Switzerland, 1, Rue des Grillons, CH- 2800, Delémont, Switzerland 2. Institut National de la Recherche Agronomique (INRA), UR 0633, Station de Zoologie Forestiére, 2163 Av. Pomme de Pin, 45075 Orléans, France Corresponding authors: Marc Kenis (m.kenis@cabi.org), Alain Roques (alain.roques@orleans.inra.fr) Academic editor: David Roy | Received 26 March 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Kenis M, Roques A (2010) Lice and Fleas (Phthiraptera and Siphonaptera). Chapter 13.4. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 833-849. doi: 10.3897/biorisk.4.65 Abstract A total of 31 Phthiraptera species alien zo Europe are listed. They include 24 chewing lice and seven suck- ing lice of 12 different families. The 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. The recent period did not show any acceleration in alien arrival in Europe. Alien fleas include six species in the families Pulicidae and Ceratophyllidae. Three of them are primarily associated with rats and are capable of transmiting major human diseases such as the bubonic plague and the murine typhus. Keywords Phthiraptera, lice, flea, Siphonaptera, alien, Europe 13.4.1. Introduction Phthiraptera (lice) and Siphonaptera (fleas) 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 flea (Ctenocephalides felis felis (Bouché)), the rat flea (Xenopsylla cheopis (Rothschild)) or the human flea (Pulex irritans L.). Although many of these are of unknown origin, they Copyright M. Kenis, A. Roques. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 834 Marc Kenis & Alain Roques / BioRisk 4(2): 833-849 (2010) are probably allochtonous in Europe, having arrived in ancient times with their hosts (Mey, 1988; Beaucournu and Launay, 1990). Thus, 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 fleas, 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 first time at the end of the 19" century, many probably came much earlier; the exact date of arrival remaining unclear in nearly all cases. 13.4.2 Phthiraptera Lice are exopterygotes* of birds and mammals. Most species are host-specific but others are rather polyphagous. They 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 (chew- ing lice), but they are presently grouped into a single order, Phthiraptera (Barker et al 2003; Price et al 2003). The 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). They include 24 chewing lice belonging to 8 different families and 7 sucking lice belonging to 4 different families (Table 13.4.1). Three of the families have no rep- resentatives in Europe (Gliricolidae, Gyropidae, Trimenoponidae; all in the Amblycera suborder). The 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 modified 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 first observed in Europe during the 18" and 19" century (18 species out of 31- 58.1%), although they probably arrived much earlier with their animal host, in most cases a domestic species. The recent period did not show any acceleration in alien arrival in Europe with only 4 species (12.9 % of the total species) newly observed dur- ing 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 farm- ing, in particular Menopon gallinae, Goniocotes gallinae and Eomenacanthus stramineus Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) 835 % species % species 60 40 20 0 0 20 40 60 a pla log ena Haematopinidae =.5™ ein i at Anoplura Echynaphthiriidae ‘edicinidae Prhiridae 4 Phthiraptera Amblycera Menoponidae G Sritok idae Ricinidu Goniodidae Philopteridae Trichodectidae Bovicoltidae 7 375 4 Ischnocera Coptopsyllidae Fiepophthalnide e gal al aed idae Ceratophlidae schnops ermi syiidae Native species Alien species 3 Siphonaptera 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. The number over each bar indicates the number of species observed per family. (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 (Moli- na)); (Laurie 1946; Newson and Holmes 1968) and the North American Trichodectes (Stachiella) octomaculatus on raccoon (Procyon lotor (L.)); (Hellenthal et al 2004). Only seven sucking lice of four families (Enderleinellidae, Hoplopleuridae, Linog- nathidae, and Polyplacidae) are considered Neozoans in Europe (Table 13.4.1). The Asian Polyplax spinulosa (spined rat louse) causes hair loss and pruritus to wild and do- mestic rats (Rattus spp.). The cryptogenic species Linognathus stenopsis and Haemodip- sus lyriocephalus parasitize goats (Capra hircus L.) and hares (Lepus europaeus Pallas), respectively. According to Durden and Musser (1994), another Haemodipsus species, H. setoni Ewing associated with Lepus spp. in North America is possibly an introduced species in Eurasia (this species has not been included here). Three species have been in- troduced to Europe with their Sciuridae host from either North America (Enderleinel- lus longiceps and Hoplopeura sciuricola with grey squirel, Sciurus carolinensis Gmelin; Britt and Molineux 1979) or Asia (Enderleinellus tamiasis with Siberian chipmunk, Tamia sibiricus (Laxmann); Beaucornu et al 2008). Solenopotes muntiacus has also been 836 Marc Kenis & Alain Roques / BioRisk 4(2): 833-849 (2010) AY» 3 G Forts é ‘\ of ~ \ iiss ie e wer ee of st es y i 0.50 mm Figure 13.4.2. Alien Phthiraptera (Anoplura). Solenopotes muntiacus female from Muntjac deer, Muntia- cus muntjak (Credit: British Museum of Natural History, London) introduced from Asia to Great Britain with muntjac deers, Muntiacus reevesi (Ogilby) (Dansie et al 1983). In addition, Haemodipsus ventricosus (Denny) which lives on rabbits (Oryctolagus cuniculus L.) can be considered as alien iv Europe, originating, as its host, from the Iberic pensinsula (Durden and Musser 1994). 13.4.3 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 fleas are listed as alien to Europe, including 5 spe- cies 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). The aliens belong equally to two families, Pulicidae and Ceratophyllidae, whereas the latter family largely dominates the native entomofauna. Three of these fleas have rats as their main host (Beaucornu and Launay, 1990). ‘The tropical rat flea, 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 Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) 837 Figure 13.4.3. Alien Phthiraptera (Amblycera). Gliricola porcelli male from guinea pig, Cavia porcellus (Credit: British Museum of Natural History, London) found sporadically in port areas and elsewhere, e.g. it was recorded from Wales in the 1950s (Hopkins and Rotschild 1953). The third species, Nosopsyllus fasciatus, is a temperate species from Asia. Rat fleas 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 Rickett- sia typhi (Beaucournu and Launay, 1990). The North American species Orchopeas howardi is found on the grey squirrel (Sciurus carolinensis), an invasive rodent in Europe (Keymer, 1983). In addition, a rabbit flea, Spilopsyllus cuniculi (Dale), can be considered as alien in Europe, probably originating with its host from the Iberian Peninsula. It has invaded a large part of Western and Central Europe (Soledad Gomez Lopez 2005). It is the principal vector of rabbit myxomatosis, a disease which was deliberately introduced from South America into Europe in 1952 in order to control rabbit populations (Beau- cournu and Launay, 1990). 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Revista Ibérica de Parasitologia vol. extra: 113-124. 843 Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) (F86T) SaqseazTS “(COOT) BANISeT Pure PAOIJIg ‘(QZ61) Wouyeypy pue son “(896 T) J2]Yory pue sousayy ‘(9961) FousaN “(FCG 1)AOYYsoTNO], ‘(6€61) vosduroyy, “(FH6I “FZ6T) (6€81 AnB9¢ “(QS8T) 198¥Iq “(S00Z) Td “IN ‘ToasTourNg) stuayy “(ZOOZ) Je 9 129194) “(€QOT) | (s#8vj0I9-) ‘TI ‘GD UA ‘Ta jorepoid snp dar01nq elyey euney ‘(1 /6]) uazinysypoig SoIepy AG Fh) Poa 6E8I orisesed snsdipomav ey seproeyddjog (2saadaad SNIVIJUNT,) BCG] ‘uosduroyy, (F661) Jossnyy Joop jorepoid SNIVIJUNUL pure usping ‘(€g6[) Je 19 aIsueq oefunyy CQ6I orisered sajodouajos | sepryyeuSoury (FS6T)Aoxysr[NoL (VGI “YC6T) (8€81 An893¢ ‘(GQOT) BYANISe'T pur PAOIZAG ‘Ja\stouIng) ‘(O88T) 298¥Iq “(6861) SOIT pur LI “UD ‘UW oyua8 | rorepord sisdouags seuouTF] ‘(€QOZ) Biya] PUNE | (~dvy) yeory 0 ae AD Ya = WO aes gcgt| -oaddr| /onisered snqqwusourTy | sepryiyeusoury I@6l (SQ0Z) Je J2 JoOUU0D, Q] SzsuUaUOLDI voroury| Jo1eperd ‘SIIIO,] VJOILANIIS ‘(6Z61) xneudjoyy pue ng SNANIIC 9/61 YVON | /onisered vanadosdozy | seprmna,doydoyzy OT6L “oyueryey (F661) Jossnyy pue SNIDAIQIS (ea1oy) | rorepoid SISVIUUD uspinq] ‘(800Z) Je 12 Nunooneag SVIULDT. al 22 orisered snpjauiapaapug | Sepi{oulsJopuy SIG6I ‘SHIA7 29 (SOOZ) [B32 JoUU0D,. O| sisuauNosvI vououry| Jo1epoid S30] [Py Ssda218u0] ‘(6Z61) xnoudjoyy pure ag SNANIIS aad YVON | /orisered snpjauiapaapug | Sepi{oulspJopuy vpanjdouy esayderryy yg SIDUDIIFIY veNqeH S311JUNOD poprauy adoing ut proses IST oSuvri DATION snjejg {Urea ca | Lapsoqgny sarood¢ PIO ‘OLOZ/€0/ZTZ aepdn ase7T *({] xtpuadde das) SIN AA 02 JoJor suoMerassqqe yeiqey ‘(| xipuodde sas) 991 ¢€ OS] 01 Jofor suOMeTADIQqe sapoo AmuNO’) ‘so1deds stuasoidA19 F adoring 01 uaITy Ww :smieig ‘odomng o7 uorye sorseds erayeuoydis pure vsaidesryiyg dy} Jo sonstajoeIeYS pur isi] “|p S| VIGUL Marc Kenis & Alain Roques / BioRisk 4(2): 833-849 (2010) 844 (suovaq ayy) AYN]; “(SnI14saULOp “(HOI ZG) ANB9g says ‘(Q00Z) PIsteuIOyN puP AOZITII J | $77]V4)) [Moj ‘(L661) OS8N pur s1AojAeg “(8T8I)| — IAsewoq vn (SI8T YoszaIN) YOSZIN “(ZS6]) IOP pue yoyonoyy | “(snuvisvg) “Ou “Td {LI ‘WA o1ua3 | r0repoid SNIUIULDAIS “(600Z) PAPI ‘(ZOOT) Je 29 1914) jueseoy gq (al. “saa eg SI8T| -oaddry] sonisered| 5 snqgjupspuauog | sepruodouayy (8961) SowyOH (snddoz CE6I pure UOSMAN ‘(9FG6[) INT “(SQOT)| —-4ezsv20d p47) vououry| ro1epeid Torey sadiloa study ‘(FO0Z) Je 29 yeEyWTOH nddovy ZG S 29D] /onisered viuanbfnyg IEPI[OSLIT]5) (86C61) AOYYSTNO]T ‘(FOOZ) Je 39 WAalo1g “(HYGL “HT6L) AN89g “(O88I) °3eId “(ZS61) PIOW Pur roypnoyy “(G00Z) | ——(sjjar4od Td ‘LI ‘AH sTUayI ‘(Z0OZ) [B19 1D “(€OOT)| 7747) ssid WH ‘Ud ‘Td ‘Sa voroury | Joyepard 8EgI Toasiouring ele] euney ‘(8Q0Z) Neepsiog eouInry “Ad “HO ‘Dd ‘LV SEsI S$ 29D] /onisered sypao sndoat) sepidorcr) (2CC6I) AOYYSIPNOL, (PEGI ‘P761) ANB9S “(T8ZT) YUIYDS ‘(O88I) 1983eIq ‘(6861) Wuzipereg ‘(ZS6T) POW pure yoyonoyy “(S00Z) (smppaasod I$ “O8"1d “Ll stuay “(ZOOZ) Je 19 199194 “(€QOT)| 7724) ssid ‘VE AS “ied “Cd voroury| Jorepoid (L8ZT yueryps) elyely] euNey “(QQOZ) [Neepiog eournry {| “Ad “HO ‘Dd ‘LV I8ZI $299] /onisered| vy 1a24od VJOILYD) IEPIOONT]S) piayquy viaidesryyg (YS6T) AOYYS?INOL (F007) Je 39 otaa0l0Ig “(FBG 1) SaqiIoaTIS “(HH6L) ANB9¢g “(COOT) PAANILT Td (6€81 pure PAIS “(G0OZ) SIU>>I (Z861) (dds WH Ua ‘Td ‘Sa rorepaid ‘Jaasjourng) Je 39 ZOWIOL ‘(ZQQZ) Je IP JNA] suzqMYy)SIeY Ee (a ao ul SI eI eIsy | /orisesed vsojnurds xodjog seproelddAjog ea adomy ur| oSsues Lapsoqny S9dUIIIJOY WIUIQGeP | Se1UNOD popeauy] | poser IsT | saeNY | oUTTSay | snIEIG saroodg¢ J2pPJO 845 Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) (FSS6T) AONYSI[MNOL, “(H8G6T) Ou BOqIoagT!S “(FH6 1) 4n389g “(C00Z) “Td “IN ‘ET SLI (I8ZI yuerys) PYANISeT PUL PAOTJIG ‘(FOQZ) |e ‘AH ‘AD SW ‘TW o1ua3| rorepoid sudo (yjos1a0g) qo yeyITPOH (9161) sSurmumn (s1acQ) daays fl) S9+79“)9-a9 9I61I| -oiddAID] sontsered) 5 yjomuuog JEPIOOTAOG DAIIOUGIST viaidesyyg (819129404 QCQT Toaistouring (L661) PIO! pure royonopy| vzav7) ssid Ta Se al voloury | Jo1epoid unpidsie sepruod “(SOOT) STUDS “(ZOOT) Je 39 JoNe+y eouIner) Wi “Ad “HD ‘IV 9961 S$ 29D] /onisered uodouauitsy -OUSUTTIT (L261 ‘eyDAZIOIOTZ 29 (snuvisvq[) jorepoid eYsSMI[IZIPOY) (SOOT) BYANAseT pure PACES) —-JURSKOY Td “AC ‘ZO 8661 jorsered ruvisogd vpigoy) | sepruodousyy (ZS61) AoxYseINOL “(HH6T) A ba lS OU “Td “UWS (Ty8T Auusqd) An893¢ ‘(OS8T) 198g ‘(600Z) PAN] ‘(09]07) al Fl ig) bi oY = ers es joyepoid UNIDUIGAN “(ZOOZ) [B19 JoMNd+y ‘(Z7HQI) AUG, suooyey] WW ‘Sd “dq ‘pq td /orsered wunjvydas0djov0ayy | Sepruodousyy (sma14sauLop “(EYL “HT6I) An89g “(S00Z) 4ASSYI) (O88 “198eIq) BYANISL'T PUL PAOTJIS ‘(088T) Mosreds TRA jorepoid vyviasvfiaponb yaselg ‘(FO0Z) [e 39 eyoTPy asno}] WA “Ad “ZO “Ag O88I orisesed vapIsy sepruodouayy (26C61) AoyYysomnoy, “(H861) BoqIoaTTIS “(PEEL “b76I) An83¢ “(TZT) YAeIYPS (9007) | (#SvajA/V) PsIeUIOy] Pue AOZITPIJ ‘(O88T) AdyINT, a1eSiq ‘(1661) 2ISaN pur s1aopavg | <(snozgsausop Wn ‘(LS61) PIOW pur iwoyonoyy “(600Z) snypvs ‘Su ‘OU “Td SLI PAST] (FOOT) Te 39 [eEWIUSTIEH | $777VD)) [oj ‘AH ‘FD WA ‘Ts jorepoid Cele D “(ZOOT) [e 39 UPD “(ZH8T) AUUEq| —IAsou0q ‘Sd “AC ‘Dd “Ad I8ZI jorsered avuyjvs uodouayy| sepruodousyy (VL61) AOYYS[NOL, “(HHI Ou (O88T 18eIg) E761) ANBIg “(OS8T) 198¥Ig ‘(600Z) he WL oa 1a o1ua3| so0repoid VID] VIJUDBID PAST] (FOOT) Je 39 [eyUeT[EH, pquinjo’) ‘oq “Ad ‘Dd “Ad Sa -oidAIZy orisered DIJA1ISAOGY OP] sepruodouayy ‘Leads | adoimy ur| oSsues Lapsoqguy S9DUIIIIOY yeIQeP | SatUNOD papeauy] | poder JsT | aATIeNY s1}e1¢ sarood¢ J2PJO Marc Kenis & Alain Roques / BioRisk 4(2): 833-849 (2010) 846 GuspapaY) AdYINT$ “(SnI14SaULOp (6Z81 (@CC6TL) AOYYSoTNOT, snus jeordon SJaquaypsvy ) “(PZ6T) An82g “(GOOZ) PAPI “(H007) | 5777%D) MO] Td LI ‘dD Wd -qns | Jorpaid sas ye 39 FeyIUATOH ‘(ZOOZ) [e 39 Jowa+y STISOWIOC] ‘oq “Ad ‘Dd “Ad ‘eotdosy, /orsered sng dvq0190Uuans depIpoluor) (SMIVISLAI oayuy) [Mojeag uvIpUy] ‘(S15 VOIIUL DpIUnyy) 8I8I “Y9SZIN (O881) 198eIg | [MoT vourNr jorepoid SNIVINSUDIIAL “(QI8 I) YOSZIIN ‘(ZOOZ) Je 9 Jouery parowyapy OW ‘NH “Ad orisered S940I01U0L) depIporuory (SWIVISLAI (86661) AoyYseNOT, oar) “(EYL “HT6L) An89g “(€007) paoyeag OU “Td ‘LI ‘AH sorepord (S¢ZI ‘snoeuur) eryeq euney ‘(7QQZ) [e 39 Jour ueIpu] MA TA “ad “Dd orsered stuoand saporuory depIporuory (86C61) AOTYSIINOL, (HHT) 489 *(900Z) TYSIeUION] PUP AOZITIIJ “(O88T)| (S#IsSsaUOp 198eIq “(166 1) PIS9N Pure dAojarg snes VN ‘Su “Id (SZLI1 “(LS61) HOW Pure wyonoyy “(FOOZ) | 577%) [oJ ‘LI ‘NH “Wd ‘TA rowepaid 199") O({) avuypys ye 39 FeyIUaTPAH ‘(ZOOT) [e 39 Joqsy STISOWIOC] ‘oq “Ad “Dd “Ad orisered SIJOIOLUOL) dePIpOluor) “(HCL “HE6T) ANB9g “(O88T) 8eIg (dds Sa PL8I PIPED “(H007) Te 39 TeywET[PH “(ZOOT) | — s#uvISGI) ‘OU “Td “IN ‘LI royepord snppgdasoshaga [e 39 Jouary “(€QOZ) Pye euney Jueseoy ‘ANH WI Ag “Ag orisered SIJOIOLUOL) depIpoluor) “(Hyol| (sve) “676 1) An89¢ ‘(CQOZ) PYANIge’T] pue Aoyn], OU ‘Ld “Td (S¢ZI ‘snevuury) PAOIJIS ‘“(O88T) aselg “(ZGG61) JPOP | Poeonsouop “TN
  • jorsered soa OPAT: || HRpEesbe denon, CIGL Sued (4010 SNIVINIVULOIIO (SOOZ) StUPy (F007) uolI0Af) jorepoid (277]21G9V4S) ye 39 FeyIUATPAH ‘(ZOOZ) [e 39 Joya WOODDEY qd “HO “Ad {Ly} usouyuy orisered SHIPPOYNAT | VePNI9pOySN], (8€81 JojsTouing) (FH6L) ANB9S ‘(088I) I99kIg snizadvaqhjod “(Z61) [IO Pure royonoypy “(ZQOT) | —_ (sz4sva/apA7) o1ua3| ro0zepoid (snanady{xQ =) fe 39 Joua+y (€QOZ) PleI euney AYINT$, LI Wd “Ad ‘Dg O88} -oIddAr_D | soratsered snanadyngyay sepraidoypryg (X21p1ag xIApla ) asprieg (FOOT) Je 29 Jey UeTPPH “(SNI1GI] 09 SC6T ‘ARID ‘(ZOOT) [& 39 JONAT “(EOOT) Pe] svuvisvg{) OW “Td ‘LI ‘NAH jorepoid SNSO]NIDUL euney ‘(Z/6T) F[oqrtq ‘(8¢6T) ARID yueseay “aD “ag ‘7D “Ad SC6I eIsy| /orisesed snanadiT seprioidoypryg (SmI149]09 6F6I ‘UOsIsUIy (SOOT) BYANIseT pue PAOIJIS ‘(FO0Z7) SNUVISDY[) festa ci joyepoid SNILYIOI fe 39 EwUITPPH “(ZOOZ) [B29 JaHEH] —_—-usvoyT Td “AC “ZO “Ad 6861 eIsy | /orysered snsaodosvT | — seproadoyryd (86S61) AOyYys2|noy, “(HHGL ‘HE61) ANB9g “(OB8I)| — (sIZssa1U0p (9981 [?q9ID 199eIq “(LC61) PFO Pure ryonoyy snypvs VN‘OU “Td “IN UT YOSZIIN) (FOOT) Te 29 FEsUETIAH “(ZOOT) | 57477VD) Joy Se Cie aa “1a o1ua3| rorepord DY Uvl50194aY Je 39 Jou9+y “(€QOZ) PyeI vuney onsouroq, ‘oq “Ad ‘Dq “ag 9/81| -oiddArZD | /oratsered Lagspsoqojny)| seprsaidoyryg (TPH8T ‘Auusd) (YOOT) | (suuwssvq[) OU “Td ‘LI royepord BG HOR fe 39 emUeTPH “(ZZ6T) BoqeId| ——_ dueseoyd ‘Sd “AC “ZO “Ad sha BIsy | /rsered vyjayoizio10jz | —_ epiporuory pee adoingq ur| + osuer Lapsoquy sadUaIafaY yeyIqey | SetIJUNOD papeayy] | prooar ysT | = aATeNy snj}e1Sg sarood¢ JIpJO Marc Kenis & Alain Roques / BioRisk 4(2): 833-849 (2010) 848 MS ‘SU ‘OU ‘AVW-Ld ‘OZV (6002) z9doT “Ld ‘Id “Td “IN ZIWIOT) PEprfOs “(996I “ZS61) WwS ‘LI “AW ‘NT ‘ET ‘(€E96T) SMAd “(800Z) TP 39 PHF seplinyy ‘OISSLI “UVSALI “(LIGT) esneryy (0061) OSTRA Joyo ‘LI “AI ‘NH “UD -IIJPD “(8661) [eoseq pue nusoonesg pue ‘dds ‘5 YOoN’A (OO8I SnUYP ‘(0661) Aeune’y] pure nusooneag sayy “dds Wd ‘Od “TVa-SA osog) sugpiasvf ‘(F86[) JOAooTy pue nusoonvag snuapody ‘oq SIG “Ad -odwia] | orepaid (sappdsdosopy) ‘(SZ6L ‘9Z6I ‘TZ61) NUJooneag “dds SNUOM ‘T) “HO “Ad ‘LV -PISy /orsered snyplsdosong sepryAydoresa) MS “OU ‘GVW “Ld ‘OZV-Ld ‘Ld “Td “IN (LW SIW GA Ohl 1a ‘OIS-LI UVS-LI ‘LI ‘SI “AI ‘NH WH “dD “Das “aD SANecd)) TUWOUD UD (seay WOO-Us UW Ing) SepLINyy ‘Od ‘Td ‘NVO-SA (F007) | = Yo pure “Tvd-Sa ‘Sa “Aa (LISI 19 1A99(016 ‘(600Z) Zado7T zaurory | sugy4 snuvY So aaeZD ‘IIYUQYIS) s7uUsas PEPITOS “(668T) PISO “(SZ8T) | ‘suaessousop XO ‘HO ‘Dd “Ad o1u98 | r0repard (xy4sdordoT) eq ‘(0661) Aeune’y] pure nusooneag Sn ‘Vd ‘LV ‘dV “IV -oidAr) | /orisered wydsdogdaT | seprfAydoresszy (asues saneu (O61 oy} Ul y1a77 ‘JJO]) uzuUas YqQuLoIO’)) jorepoid (ppp sdouzuar)) (C861) UURUTTYDsoY puke NUJOONeIg ueyy eIsy| /orisered mypdsdoyvy | sepy[dydoresa) vioideuoydis | Lhida | asurs Lapsoqugy SIIUIIIFOY WIUIQey | SersyUNOS popeauy] | poser 3sT | aaTIeNY snje3¢ sarood¢ PIO 849 Chapter 13.4: Lice and Fleas (Phthiraptera and Siphonaptera) “(786 1) Je 19 sowrey-ororede7z ‘(POGI) TyPsoqesT], “(EZGT) UOueT, ‘(BZ61) ANBIg “(7HG6T) PIofeN JO 101D9A “SHNISNUL SANT NU ‘Ld “GVW “Ld ‘OZViLd “Td “(1Z61) FART “(ETOT) O1UPAT] ‘suewny LW LE DISsLI (€O6I ACOG1) STO S6 1 pOpeyo DT) | ey Hee UE “al ‘NH “UD “ad PESO) pure eyjoises euvies “(Q661) Keune | — ‘smo75aaL0u WOON WA JIN | Joreposd sidoaya sidoaya pure nusoonvag ‘(/761) [e 9 preusog SHEE ‘NVO°Sd ‘SH “Ad YOGI yeoupy | /opsered vypdsdouax seprornd snyd ouTINu “(ZO61) HWS “(TH6) BIAeN | pure onsejd (FOG6L IOyxeq) “(€C61) P[Eyosioy, pur surydopy JO OID9A jorepoid SISUAYIISVAG ‘(0661) AeuneyT] pue nusooneag| “dds suzy gD ‘NVO-SA THI eolyy | /onisered wyplsdouay seploiyng snjnIIUNI SNBVIOJMUC) ‘snuppisoyf (€O07Z)| — ssupzaj5 (FO6T Txeq) elyeq] euNe ‘(1 86[) Je 39 NuIJOONvag uqqes vououry| Jo1epeid stuffy syoi9y8 ‘(ZZ61) Aeuney] pur nusoonesg | = ‘JreU07109 LLOI YON | /orisered snpydsdojdognq seproyng sSnNIIUNI SNBVIOIMUC) ‘sadjna sadjn ‘DUP DULY ‘Sy8 SYD) ‘SNJOILUID shuou ey sae <(jorzmmbs Ao18) C681 (SZ6I)| sesuautposva voloury| Jo1epoid iayeg ipivmog adioy stud ‘(f66 |) snowAuouy SNANIIC OO8I YMON | /orisered svadogo4c | zepy[dydoresa) [ee adoinq ur} + osuer Lapsoqguy SdDUIIIFOY WIUIQep | SersyUNOD popeauy] | poser ysT | saTIeNY snje3¢ sarood¢ PIO