BioRisk 4(2): 793-805 (20 l 0) Apeer-rev ‘iewed open-access journa 1] Sohaaee eee: BioRis www.pensoftonline.net/biorisk Psocids (Psocoptera) Chapter 13.2 Nico Schneider 79, rue Tony Dutreux, L-1429 Luxembourg-Bonnevoie, Luxemburg Corresponding author: Nico Schneider (nico.schneider@education.|lu) Academic editor: David Roy | Received 1 January 2010 | Accepted 25 May 2010 | Published 6 July 2010 Citation: Schneider N (2010) Psocids (Psocoptera). Chapter 13.2. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(2): 793-805. doi: 10.3897/biorisk.4.46 Abstract Among the 231 species of Psocoptera present in Europe, 49 (21.2%) are considered to be of alien origin. They include 29 exotic introduced species and 20 cryptogenic species. Most of the exotic species origi- nated from tropical and subtropical areas, essentially from Africa. Many of them are food pests, moving along with stored products. Thirty-nine of these species occur in buildings in Europe. Keywords Psocoptera, psocids, domestic, stored products, alien, Europe 13.2.1 Introduction Psocoptera (commonly called psocids) are one of the smaller orders of paraneopteran insects. Many species are arboreal, but a few are more usually found on low vegetation or in litter. All feed on microflora and organic debris. Some are found in nests of birds and mammals, within aggregations of other insects or associated with human habita- tions. The head of these usually soft bodied pterygote insects (with a body length of 0.67 mm to 8 mm) is globulous with an usually prominent clypeus and projecting eyes, long and filiform antennae and biting mouthparts, the laciniae being characteris- tic for the order. Adults have usually four wings with simple venation. However, many species are brachypterous, micropterous or apterous (Lienhard 1998, Lienhard and Smithers 2002, Mockford 1993, New 2005). Copyright Nico Schneider. 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. 794 Nico Schneider / BioRisk 4(2): 793-805 (2010) 13.2.2 Taxonomy of Psocoptera species alien to Europe According to Lienhard (1998) (Lienhard 1998) a total of 231 species in 25 families of pso- copterans are present in Europe. Forty-nine of these are not native, 29 as aliens to Europe and 20 as cryptogenic, globally accounting for 21,2% of the total psocopteran fauna in Europe (Figure 13.2.1). They are included in 12 families: Lepidopsocidae (5), Trogiidae (4), Psoquillidae (3) and Psyllipsocidae (5) belonging to the suborder Trogiomorpha; Lipo- scelididae (14) and Pachytroctidae (2) belonging to the suborder Troctomorpha; and Cae- ciliusidae (1), Ectopsocidae (10), Elipsocidae (1), Lachesillidae (2), Peripsocidae (1) and Psocidae (1) belonging to the suborder Psocomorpha. Details for each family are as follows. Lepidopsocidae: Adult wings of lepidopsocids, which belong to the family group Atropetae, are generally pointed apically when fully developed. Body and forewings are generally covered with scales, but occasionally with dense setae Mockford 1993. The five species found in Europe are all alien. Trogiidae: Species in this family, which belongs to the family group Atropetae, are apterous, micropterous or brachelytropterous. Body and forewings lack scales and dense setae. Four of the 19 species found in Europe are cryptogenic (21%). These four species are brachelytropterous, presenting short, leathery winglets similar to short elytra. Their habitats are associated with humans, e.g. within buildings and glasshouses (Lienhard 1998, Mockford 1993). Psoquillidae: Body and forewings of species in this family, which also belongs to the family group Atropetae, do not bear scales. The three species currently found in Europe are not believed to be native, two of them being with certainty of alien origin and the third cryptogenic. All occur within buildings in Europe. Psyllipsocidae: This family belongs to the family group Psocatropetae. The five species found in Europe (100%) are either alien (Baz 1990) or cryptogenic species (Baz 1988). All are usually found in buildings and glasshouses in Europe. Liposcelididae: ‘These psocids belong to the family group Nanopsocetae. They are characterised by a flattened body and antennae with secondary annulations on their flagellum. Fourteen species out of the 39 (36%) found in Europe are either aliens (Broadhead 1950) or cryptogenic species (Broadhead 1954b). They include exclusively apterous species, all of which being occur in buildings. Pachytroctidae: The body shape of the species of this family, which belongs to the family group Nanopsocetae, is not flattened dorsoventrally and the basal flagellar seg- ments are not secondarily annulated (New 2005). Only three species live in Europe, and two of them are not natives (67%). Caeciliusidae: This family belongs to the family group Caeciliusetae and to the superfamily Caecilioidea. The species are characterised by the presence of at least one, or more commonly two or three, ventral abdominal vesicles (Mockford 1993). The family was once named Caeciliidae, but the latter name was changed because of ho- monymy with a family of amphibians (Lienhard 1998). Only one species out of the 15 (7%) found in Europe is cryptogenic. Lacroixiella martini, is only known by the two syntypes collected by Lacroix in 1918 in a french military hospital (Lacroix 1919). Psocids (Psocoptera). Chapter 13.2 795 7 species % species 40 30 20 10 0 0 10 20 30 40 eS ee 25 Mi Liposcelididae 4 H- _ Ectopsocidae 0 | Lepidopsocidae 0; Psyllipsocidae S=_ rogiidae 0+ Psoquillidae Native species 0 == Lachesillidae Alien species | | Pachytroctidae 14mm Caeciliusidae 3 Elipsocidae 8 mi __— Peripsocidae 3 Psocidae 2 ~ Amphientomidae 2 # Amphipsocidae | | — Asiopsocidae i Epipsocidae 2| M- Mesopsocidae t+ Myopsocidae 3 # _ Philotarsidae 2 | _Prionoglarididae 3 g- Protroctopsocidae 2 g. Pseudocaeciliidae | |. Sphaeropsocidae 4 ™ Stenopsocidae 7 m _ Trichopsocidae ooo oo o 0 oo 0 00 OOS Figure 13.2.1. Taxonomic overview of the Psocoptera species alien to Europe compared to the native fau- na. Species alien to Europe include cryptogenic species. Families are listed in a decreasing order based on the number of alien species. The number over each bar indicates the number of species observed per family. Lachesillidae: These psocids belong to the family groupe Homilopsocidea. The species have an areola postica* in their forewings characterized by a very sparse and short ciliation on its veins and margin. The /acinial* tip of the Lachesillidae is slender and bicuspid *(Lienhard 1998, Mockford 1993). Two species out of the 12 found in Europe are aliens (17%). Ectopsocidae: Members of this family, which belongs to the family group Homi- lopsocidea, are characterised by the absence of an areola postica in their wings such as in the family Peripsocidae. Ten out of the 14 species found in Europe (71%) are either alien (Broadhead 1950) or cryptogenic species (Baz 1988). Six of these species are found occurring within buildings, either regularly (Baz 1990) or sometimes (Baz 1990), compared to only one of the 4 native species. Peripsocidae: Species of this family, which also belongs to the family group Homi- lopsocidea, are also characterised by their absence of an areola postica in their wings. Peripsocus bivari is the only alien among nine species found in Europe (11%). 796 Nico Schneider / BioRisk 4(2): 793-805 (2010) SS ae Cryptogenic Tropical/subtropical > 408 14.3 Central & South America _ 4.| North America. 6.| Asia-Tropical Australasia 2-0 4.| ) Africa, Asia Africa Asia 6.1 18.4 4.| Figure 13.2.2. Geographic origin of the Psocoptera species alien to Europe. Numbers indicate the rela- tive proportion of alien species originating from a given region. Elipsocidae: Veins and wing margins of species in this family, which also belongs to the family group Homilopsocidea, bear setae but the hindwing marginal setae are limited to the radial fork (New 2005). Propsocus pulchripennis, occurring out-of-doors in France, Great Britain and Madeira, is the only alien among 24 species found in Europe (4%). The native range of this widespread species is the coastal regions of sub- tropical countries (Lienhard 1998, Lienhard and Smithers 2002). Psocidae: This family belongs to the family group Psocetae. Its members are easily recognised by their wing venation, where the areola postica is fused to the M-vein. The Psocidae fauna of Europe includes 34 species but Trichadenotecnum innuptum is the only alien among them (3%). 13.2.3 Temporal trends of introduction in Europe of alien psocids The cryptogenic species Trogium pulsatorium was already known as Termes pulsatorium in the 10th edition of Systema Naturae by Linnaeus in 1758 (Linnaeus 1958). From 1850 to 1874, three other cryptogenic species and the alien Psoquilla marginepunctata were recorded from Europe. One alien and one cryptogenic species followed in 1899. In the 20th century, seven species on our list were recorded for the first time in Europe from 1900 to 1924, 18 from 1925 to 1949, seven from 1950 to 1974 and finally ten from 1975 to 1999. In the 21st century, no new alien has arrived in Europe to date. 13.2.4 Biogeographic patterns of the Psocoptera species alien to Europe The distribution status is only known for 29 species out of 49, 40.8% being thus considered as cryptogenic. Figure 13.2.2 details the probable regions of origin. Most Psocids (Psocoptera). Chapter 13.2 797 20°W 0° 20°E 5o°N= 30°N= Number of alien species no data 1.0) 1s 20-31 Figure 13.2.3. Colonization of continental European countries and main European islands by the Pso- coptera species alien zo Europe. Archipelago: | Azores 2 Madeira 3 Canary islands. species appear to originate from Africa (9 species, 18.4 %), far beyond the other conti- nents but for seven species (14.3%) we only know that they arrived from tropical and subtropical regions. Central and Western Europe appear to be more colonized by alien psocids (Figure 13.2.3). 13.2.5 Pathways of introduction in Europe of alien psocids, invaded habi- tats and known impacts The main pathway of introduction is trade. Lachesilla pacifica is probably dispersed by wind. Most aliens and cryptogenic species are found in warehouses in stored products. Many of them are food pests. Forty species (88.9 %) are associated with buildings in Europe. 798 Nico Schneider / BioRisk 4(2): 793-805 (2010) a Figure 13.2.4. Alien psocids. a Ectopsocus briggsi McLachlan, 1899 (Credit: Tom Murray 2008) b Lipos- celis bostrychophila Badonnel, 1931 (Credit: Joyce Gross 2006). Acknowledgments I thank Dr. Charles Lienhard (Muséum @histoire naturelle de Genéve, Switzerland), and Drs. Alain Roques and David Lees (INRA, Centre de Recherche d’Orléans, France). Ms. Joyce Gross (BSCIT, University of California, Berkeley, USA) and Mr. Tom Murray (Groton, Massachusetts, USA). 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Selys-Longchamps E de (1872) Notes on two new genera of Psocidae. Entomologists Monthly Magazine 9: 145-146. Titschack E (1930) Die Copeognatha, Megaloptera, Neuroptera und Mecoptera der naheren und weiteren Umgebung Hamburgs. Verhandlungen des Vereins fiir Naturwissenschaftliche Heimatforschung 21: 104-127. 801 Psocids (Psocoptera). Chapter 13.2 (ZOOZ) SHUG pue preyusrT “(866]) Prequel] >) Ui ‘HO} HO ‘9861 (600Z)-(Z00Z) PreyUATT “(Z00Z) sz9yITUUS pue preyuery ‘(8661) PreyyerT “(Z8G6T) 081g “(066I1) ZA x avw CAVW:Ld “dD “Us| «Ld “1861 POLIOUY YON V jeordongqns qestdo1y.}| ov O€6I ‘uewdeyD voifiavd vyyIsagIVT PEP Soyer (6681 ‘SUP }ed) stuuadiagynd snsosdorq. 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uMOUyUN) suaosaqnd syaIsoWtT S) (ZOOT) SFoyAUIS pure preyuarT “(OTOT POSSIN OT UT srver O66T “PreyuerT aN ~7O0Z) PIEYUATT “(866T) PHEYUATT ‘(OS6T) Peeypeorg Le YH “AD “UA ‘Td ‘Sa “AC “ZO SHO ‘LV ee suvutdvad syaosodT 8 (ZOOZ) StoyIWUS OSGI ‘peeyproig SS pur prequary “($661) PreqUrT “(OS6T) Peeypeorg is) CVWGLd ‘LI “aD ‘NVO-Sd}| FD ‘SHI uMouxuy) yynsavd syeosody] S (C¥61) & ueurread “(6007)—(Z00Z) PreYUPrT “(Z00Z) stoyITUUS CHL aS pure preyusry ‘(g661) prequery] ‘(OS61) peeypeoig { OA SLI WH SAD ‘SA ‘ZO SAT! «6D ‘OFGT umouyuy) ‘ueuseag yyavd stjassod1T X { dD} dD ‘Pc6l eorfyé| V PANISGO SYAISOGLT Bur PIey UT (66D) PEeOUIEL (eycG |) peoupeorg (9¥61) uewrsead “(6007)-(Z00Z) PreyuarT “(Z00Z) ssoyATUS pue preyuery “(866T) PrequarT ‘(OS61) peoypeosg (6761) ueurread “(6007)—(Z00Z) PreYYPIT “(ZOOT) stOYIUUS Pue PIeyoory (R66 DL) PreyuorT “(OSG 1) peeupeoig NA dD ‘LI ‘UH “AD “Us SNVO-Sd ‘SA SHO] “Ud “9F6T 9¥61 ‘uRUIIeag xvpuaul s11a9s0d1T (ZO6T “Ule}9pug) 6) wigdowoqua siassodlT NA ‘OZV-Ld ‘Id ‘LI {} “TIL UH ‘AD ‘TH ‘Sa “AC “ZO ‘AO SHO] =D “6761 uMouxU) NA “AS ‘OU ‘AVW-Ld ‘Ld “Td ‘ON “IN ‘LW SMW ‘AT ‘NT SLI (S76) UeUTIeAY “(ZOOT) SHOYTUUIS “TI ‘NH ‘UH “UD “dD “Wd ‘Td “NVO (S761 pue preyuery “(866T) PrequarT “(OS61) Peaypeosg {| -SH ‘SA “Ad “AC ‘ZO ‘AO ‘HO “AG LV] AD ‘SOI OUR. “URULTRDY) 40/09ap sya9s0d1T satads SaUDIIJOY ywrqey Sd1IJUNOD poprauy pyoser IST oSuvs saneN snyej¢ Ajrure.y Nico Schneider / BioRisk 4(2): 793-805 (2010) 804 NA “AS “NY ‘OU “AVW-Ld “OZV-Ld “Td “ON “IN ‘AT ‘LT CLI ‘SI “TI “aI ‘QH “YH “UD ‘aD “Ud ‘Id ‘NVO-Sd ‘Sd “Ad “Ad AGS ZIMA IS Ae Ver Ty: NA “AS ‘NY “OU “Td ‘OZV-Ld ‘Ld “IN “MW ‘NT ‘LI ‘SI “TI ‘NH “UH UD “AD “Ud ‘Td “NVO-SA ‘Sd “MC “AC “ZO ‘AD SHO “Ad “LV dS ‘OZV-Ld “Td “ON ‘NTI “aI “€D “Ud ‘Td “AC °ZD “HO “dd ‘LV NA “AS ‘NY “OU “AVW-Ld ‘OZV Ld “Td “ON “IN ‘NT ‘LI ‘ST ‘NH UH UD “AD “Ud ‘Td “NVO-SA “TVA “Sd ‘Sd “MC “AC “ZO “HO “dd ‘LV (ZOOT) SHOWS puke preyuerT “(8G6T) Preyusry] (ZO0Z) sIoyrUS pure preyusry (8661) prequeryT ‘(COG1) Ulspepuy (RT E6]) uewesg “(ZOOT) SI9yIWUS pure preyUarT “(866T) PIPYUTT Ad ‘S061 ons { dD ‘O€61 (ZOOZ) sOyINUUIS pur preyuory “(8661) Preyuery “(OS81) uepdsy ad ‘Ossi NA “AS ‘NY “OU “AVN “Ld ‘OZV-Ld ‘Ld “Td ‘ON “IN ‘NT ‘LI “TI “al ‘NH “YH “UD “dD “Ud ‘TA ‘NVO°Sd ‘Sd “AC “ZO “HO “Ad ‘LV (ZZ81) sdureyosuoT -s4]9¢ “(Z7007) sroyug pure preyusry “(TOT -ZO0T 8661) PreyuerT (O6I) Beeqry “(ZOOT) SJOyNWIG pue preyusTyT ‘(QG661) PreyusrT (O€6T) AYSSUL, “(€661) PIOPPOW “(ZOOT) SHoyUIg pue preyusTy] ‘(866[) preyusryT (ZOOZ) SfoyIUWUIS pur preyusTT “(€661) Japlouyss pure preyusry “(O107 -Z007) prequery (Et UA ‘ZZ81 {‘H CAVW-Ld “LI “TI ‘NVO-Sa}| LI ‘6681 LI ‘Ud “Sa “Ad *ZO ‘HO “Ad TV) — LI ‘Z061 IN ‘NT ‘LI “AI ‘SA ‘SHO “Ad| =NT ‘8861 AASS AS: “Td "ONE TT “TI “Al ‘NH ‘YH “AD “Ud ‘Td “NVO-SA (ZO0Z) sHoyINWUS pur preyuery “(0107 -ZO0Z ‘8661 ‘ZZ61) PIeyUar] HO ‘€Zol ‘Sd “MC “AC “ZO “HO “Ad ‘Va ULV (6761) UUreINg “(ZOOZ) sI9yIWUS pur preyuerT ‘(8661) PIeyUsTT BE “(ZOOT) SH9yIWS pue preyUaTT ‘(8G6G61) preyuery ALL SIDVIIIFIY yeuIgeuy SoIJUNOD popeauy PlOdo5 IST uMmouxuy) ) umouxyuy) uMouxyul) umouxuy) ©) umouxyu) jeordonqns ‘jestdory, umouxuy) uMmouxuy) ewouyy| WV uMouxyus) eS 7 TN adoimmy UT aSurs saneN snqejsg (8SZI ‘snovuurT) unisojvsnd WUNtSOL] SO6T ‘UP Popuy snqpyngiqas snjourdaT Tol ‘ueurreag stanuqd snqourdaT OS8I ‘Uap uoa snuyinbut snyourdaT sepnisosy, ZTZ8I ‘sdureyosuoT -sh]9¢ wanguivs snoosdiy sg 6681 ‘eseqry tunjgoy] sodoLgqvIOSy €88] ‘UoIEy vpyyod xtsaqddoq T66T ‘sdoyaTUS stuuadisuoy xtsagddsoq (8S6I “stetprurg) voysauop xtsaqgtsoq sepposdiyypAsg (GCG “Ueutea) snutsdasad snoosdokyy (1E6] “Ueuea) syiavdsip snoosdokagy satads Ajrure.y Psocids (Psocoptera). Chapter 13.2 805 Table 13.2.2. List and characteristics of the Psocoptera species alien in Europe. Country codes abbrevia- tions refer to ISO 3166 (see appendix I). Habitat abbreviations refer to EUNIS (see appendix II). Last update 31/ 12/ 200 Family Native range | Invaded countries | Habitat | References Species in Europe Caeciliusidae Enderleinella obsoleta Central Europe | BE, BG, DK, FI, G Lienhard (1998), Lienhard (Stephens, 1836) FR, GB, IE, IT, LU, ‘ ME, MK, NL, NO, and Smithers (2002) RO, RU, SE Ectopsocidae Ectopsocus vachoni Mediterranean CH, GB G, J Lienhard (1998), Lienhard Badonnel, 1945 region and Smithers (2002) Lachesillidae Lachesilla greeni Central Europe | BE, ES, FR, GB, IT,| G, H, J | Lienhard (1998), Lienhard (Pearman, 1933) PT, PT-AZO, PT- . MAD, RO and Smithers (2002), Lienhard (2002)—(2009) Liposcelididae Liposcelis rufa Mediterranean CH, GB, PL G, J Lienhard (1998), Lienhard Broadhead, 1950 region and Sthithersi2002) Peripsocidae Peripsocus milleri Atlantic coast of | IT, YU G, J Lienhard (1998), Lienhard (Tillyard, 1923) Europe and Smithers (2002) Peripsocus parvulus Central Europe | BE, ES, FI, FR, G Lienhard (1998), Lienhard Kolbe, 1880 GB, GR, HR, IL, . LU. NL. RO. RU, and Smithers (2002) SE; YU Trichopsocidae Trichopsocus clarus Mediterranean CHG: IDE Bis es Lienhard (1998), Lienhard (Banks, 1908) region GB, HU, IE, LT, ; NL, PL, RU. SE and Smithers (2002) Trichopsocus dalii Mediterranean AT, BE, CH, CZ, G Lienhard (1998), Lienhard (McLachlan, 1867) region DE, GB, HU, LU, ied Srasetene (20 02) PL, RU Trogiidae Cerobasis annulata Mediterranean Aly BE MCE CZ, G,J,X | Lienhard (1998), Lienhard (Hagen, 1865) region DE, GB, LU, NL, : NO, PL, RU and Smithers (2002)