Apeer-reviewed open-access journa 1] BioRisk 4(1): 81-96 (2010) F . doi: 10.3897/biorisk.4.54 RESEARCH ARTICLE B | O R IS www.pensoftonline.net/biorisk Alien terrestrial crustaceans (lsopods and Amphipods) Chapter 7.1 Pierre-Olivier Cochard', Ferenc Vilisics?, Emmanuel Sechet? | 113 Grande rue Saint-Michel, 31400 Toulouse, France 2 Szent Istuin University, Faculty of Veterinary Sciences, Institute for Biology, H-1077, Budapest, Rottenbiller str. 50., Hungary 3 20 rue de la Résistance, 49125 Cheffes, France Corresponding authors: Pierre-Olivier Cochard (pierre-olivier.cochard@wanadoo.fr), Ferenc Vilisics (vilisics. ferenc@gmail.com), Emmanuel Sechet (e-sechet@wanadoo.fr) Academic editor: Alain Roques | Received 28 January 2009 | Accepted 20 May 2010 | Published 6 July 2010 Citation: Cochard P-O et al. (2010) Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1. In: Roques A et al. (Eds) Alien terrestrial arthropods of Europe. BioRisk 4(1): 81-96. doi: 10.3897/biorisk.4.54 Abstract A total of 17 terrestrial crustacean species aliens to Europe of which 13 isopods (woodlice) and 4 amphi- pods (lawn shrimps) have established on the continent. In addition, 21 species native to Europe were introduced in a European region to which they are not native. The establishment of alien crustacean species in Europe slowly increased during the 20" century without any marked changes during the recent decades. Almost all species alien to Europe originate from sub-tropical or tropical areas. Most of the initial introductions were recorded in greenhouses, botanical gardens and urban parks, probably associated with passive transport of soil, plants or compost. Alien woodlice are still confined to urban habitats. Natural habitats have only been colonized by three amphipod species in the family Talitridae. Keywords Woodlice, lawnshrimps, Europe, alien 7.1.1. Introduction The orders in the arthropod subphylum Crustacea are mainly composed of aquatic- living species, at least during part of their life-cycle. Most alien terrestrial crustaceans belong to the order Isopoda, suborder Oniscidea, commonly named woodlice. But Copyright P-O. Cochard et al. 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. 82 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81—96 (2010) several species recorded in Europe belong to the order Amphipoda, and are commonly known as “lawn shrimps” or “landhoppers”. In 2004, the total number of valid Isopod species worldwide was 3637 (Schmal- fuss 2003). Woodlice are adapted to various terrestrial environments from sea shores to deserts and have established on all continents. As decomposers of organic plant material, isopods play an important role in ecosystems (Holthuis et al. 1987, Zimmer 2002). Most European species prefer humid and moist micro-habitats (Vandel 1960) like soil, leaf litter, mosses and decaying wood. Several species are known for their myrmecophylic nature. Amphipods are generally marine or limnicolous, and only a few species can live permanently on land (mainly in the family Talitridae). Some live near the sea, on beaches where they hide under logs and dead algae and vegetation. ‘The true terrestrial amphipods live on the surface of mulch and moist ground (Fasulo 2008). Many of the habitat features of terrestrial amphipods are similar to those of isopods. These little ani- mals are most commonly noticed by their strong, rapid jumps upon being disturbed. 7.1.2.Taxonomy of alien terrestrial crustaceans Thirty-eight species belonging to ten different families were recorded during this study. The four most commonly represented families (all belonging to Isopoda) are Tricho- niscidae (seven species), Porcellionidae (five species), Philosciidae and Armadillidiidae, both with five species (Figure 7.1.1.). Two main categories were considered: e Aliens to Europe, including 17 crustacean species originating from other conti- nents (Table 7.1.1). e Aliens in Europe, represented by 21 crustacean species native to a region of Europe but introduced in another European region to which they are not native. Several other species considered as cryptogenic or cosmopolitan are probably also aliens in some parts of Europe. However, in most cases it was not possible to distinguish their alien range from the native one. Below only those species we classify as aliens in Europe: Armadillidiidae: Armadillidium assimile Budde-Lund, 1879, Armadillidium kos- suthi Arcangeli, 1929, Armadillidium nasatum Budde-Lund, 1885, Armadillidium vul- gare (Latreille, 1804); - QOniscidae: Oniscus asellus Linnaeus, 1758; - Philosciidae: Chaetophiloscia cellaria (Dollfus, 1884); - Platyarthridae: Platyarthrus schoblii Budde-Lund, 1885; - Porcellionidae: Porcellio dilatatus Brandt, 1833, Porcellio laevis Latreille, 1804, Por- cellio scaber Latreille, 1804, Porcellionides pruinosus (Brandt, 1833), Proporcellio vulcanius Verhoeff, 1908; - Schiziidae: Paraschizidium coeculum (Silvestri, 1897); Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 83 No. species Philosciidae Armadillidae Trachelipodidae Trichoniscidae Platyarthridae Styloniscidae Agnaridae Porcellionidae Oniscidae Schiziidae DI Alien to Europe |_| Alien in Europe Isopoda Amphipoda Talitridae Figure 7.1.1. Taxonomic overview of the Isopoda and Amphipoda species alien to and Alien im Europe. - Styloniscidae: Cordioniscus stebbingi (Patience, 1907); - ‘Trachelipidae: Agabiformius lentus (Budde-Lund, 1885); - ‘Trichoniscidae: Androniscus dentiger Verhoeff, 1908, Buddelundiella cataractae Ver- hoeff, 1930, Haplophthalmus danicus Budde-Lund, 1880, Metatrichoniscoides ley- digi (Weber, 1880), Trichoniscus provisorius Racovitza, 1908, Trichoniscus pusillus Brandt, 1833. Some of the species above have proved to be very successful colonizers and are currently considered as part of the native fauna in parts of Europe, e.g. in Hungary. However, their synanthropic nature and their extremely wide distribution range sug- gest a long colonisation history as it is the case for Armadillidium vulgare. In the remainder of this chapter, we will focus mainly on the species alien to Europe. 7.1.3. Temporal trends of introduction in Europe of alien terrestrial crus- taceans The total number of crustaceans alien zo Europe has slowly increased during the 20° and the early 21* centuries, but without any acceleration in the rate of arrival. Two alien species were first discovered in Europe in the 19% century, about nine species in the first half of the 20° century and only five species since then. The majority of these alien species have been found in several other countries after their discovery in Europe. However, the number of occupied countries over time has grown steadily rather than exhibiting exponential growth. A similar pattern is apparent for woodlice species alien to Europe. However, be- cause of sparcer information on this group, the date for the first introduction is roughly known for only approximately 50% of species. To our knowledge, at least six species of 84 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81—96 (2010) woodlice classified as aliens of Europe were noticed in the first half of the 20" century and only five more species since then. Thus, unlike many other invertebrate phyla, the temporal trend in alien crusta- ceans (both intra-European and alien) has shown no marked changes during recent decades. As “silent invaders” (Hornung et al. 2007) no terrestrial crustaceans are classi- fied as pests in Europe; they are elusive animals. We suspect frequently a large gap be- tween the date of introduction and “discovery” of alien woodlice species. For example, during an intense eight year survey of the isopod fauna in a large region representing 15% of Hungary, three new alien species for this country were found (Farkas 2007). To conclude, the atypically gradual trend in the number of alien terrestrial Crus- tacea in Europe could be an artefact of incomplete knowledge. Because of both the increasing worldwide trade in ornamental plants and the general ecology of terrestrial crustaceans (i.e. often hidden in soils), it is more realistic to expect a future exponential increase in the number of alien species (especially intra-European aliens). 7.1.4. Biogeographic patterns of the alien Crustaceans 7.1.4.1. Origin of the alien species Species alien to Europe almost all originate from sub-tropical or tropical areas (Ta- ble 7.1.1.). Only one species — Protracheoniscus major (Dollfus, 1903)- is likely to be native from Central Asia. For several species, their ranges are poorly known (they are also often introduced in other tropical areas). However, several species do have a precise origin. The most widely distributed alien woodlouse in Europe is the tropical Ameri- can Trichorhina tomentosa (Budde-Lund, 1893), while the most widely distributed am- phipod is Talitroides alluaudi Chrevreux, 1901. It should be noted that a least six of the seventeen alien species were originally described from Europe (Great Britain, France and Germany) after their introduction. The crustaceans alien iv Europe generally originate from the Mediterranean basin (seven species), from western and south-western Europe (five species). 7.1.4.2 Distribution of the alien species in Europe Within Europe, Crustaceans of alien origin have mainly been recorded in western countries, where they appeared first. The four countries with most species are Great Britain (11 species), the Netherlands (10 species) and Germany (nine species) (Figure 7.1.2). Comparatively few alien species have been recorded in central and eastern Eu- rope to date (e.g. only four species in Hungary). In this part of Europe, the Central- Asian P major is one of the most widespread alien crustaceans. The high number of aliens in western European countries may be linked to the high number of scientists and the intensity of soil research (Hornung 2009). Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 85 Number of alien species | nodata J -s[ 6-10 1 - 20 Figure 7.1.2. Colonization of continental European countries and main European islands by myriapod species alien to Europe. Archipelago: | Azores 2 Madeira 3 Canary islands. There are only very few records of alien crustaceans on European islands. Tricho- niscus pusillus has been reported from the Azores and Madeira, T’ provisorius and A. assimile from the Azores but these species are native of Continental Europe. To our knowledge, the only alien aliens recorded on islands are talitrids, Arcitalitrus dorrieni (Hunt, 1925) in Scilly and Guernsey, Talitroides topitotum (Burt, 1934) in the Azores and Madeira, and 7) alluaudi in the Azores and the Canaries. All these species oc- cur outdoors and are therefore considered as naturalised. The rarity of alien terrestrial crustaceans on European islands is likely to be due to the primarily introduction route being major greenhouses in large metropolitan cities (see below). Crustaceans classified as aliens of Europe are typically species which have expanded their range approximately northwards and eastwards. ‘The eastern and central countries have a higher number of these species than more westerly countries of Europe. For example, Germany and the Czech Republic, taken together, have nine species of alien woodlice of European origin, about 45% of the total in this category. 86 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81—96 (2010) Figure 7.1.3. Alien terrestrial crustaceans. a Trichorhina tomentosa (Isopoda, woodlice) (credit: Vassily Zakhartchenko) b Arcitalitrus dorrieni (Amphipoda, lawn shrimp) (Credit: John I. Spicer). A striking example of successful colonization and establishment of such species is given by A. nasatum. This woodlouse is believed to be native to Italy, southern France and Spain (Vandel 1962). Since the start of the 20" century, it has been introduced into greenhouses in a number of additional countries of Northern and Central Eu- rope (e.g. Denmark, Finland, Germany, Hungary, Poland, Slovakia, Sweden), making this species one of the most widely distributed alien woodlice of Europe. Moreover, numerous reports highlight the successful establishment of outdoor populations in several western and central European countries (e.g. the Netherlands, Czech Republic, Romania, Slovenia) (Berg et al. 2008, Giurginca 2006, Navratil 2007, Vilisics and Lapanje 2005). Some of the aliens of Europe have also invaded other continents and can be considered as very successful invaders. The most notable ones are A. vulgare, P scaber and P pruinosus. Armadillidium vulgare and P. pruinosus are probably native from Mediterranean regions. In northern temperate parts of Europe, these spe- cies are restricted to synanthropic habitats (e.g. gardens, cellars, compost heaps). P. pruinosus is one of the woodlice that has been spread most by man across the world (Vandel 1962) and can now be considered as “synanthropically cosmopoli- tan” (Schmalfuss 2003). A consequence of the dominance of Mediterranean origin for species classified as aliens of Europe is their decreasing number towards the north of the continent (Vili- sics et al. 2007). In the northernmost countries of Europe (e.g. Finland (Vilisics and Terhivuo 2009)) only the most tolerant habitat-generalists, as well as intra-European aliens, are able to become successfully established. Alien terrestrial crustaceans (Isopods and Amphipods). Chapter 7.1 87 7.1.5. Pathways of introduction of alien terrestrial Crustaceans Because a great majority of the first isopod introductions were recorded in greenhous- es, botanical gardens or urban parks, it is clear that many were associated with passive transport of soil, plants or compost. With few visible effects in such biotopes, terrestrial crustaceans colonize and spread as undetected “silent invaders” (Hornung et al. 2007). Thus, most introductions were unintentional. The one known exception is the spread- ing of 7’ tomentosa, commonly sold as pet food, triggered by trading activity in Europe. This probably explains why, among all the alien crustaceans, 7’ tomentosa is the most widespread species in Europe. Another interesting case is the Mediterranean species P schoblii. This myrmeco- phylous woodlouse is a commensal of the ant Lasius neglectus Van Loon, Boomsma & Andrasfalvy, 1990 and was first recorded in Hungary in 2001, a few years after the introduction of the ant. P schoblii was probably introduced at the same time as its ant host (Tartally et al. 2004). It has since been found regularly (Hornung et al. 2005, Tar- tally et al. 2004, Vilisics 2007, Vilisics et al. 2007) and is now considered established, as is L. neglectus. 7.1.6 Ecosystems and habitats invaded in Europe by alien terrestrial Crustaceans To our knowledge, the only alien crustaceans invading natural habitats are three talitrid species. Arcitalitrus dorrieni has invaded leaf litter understoreys of deciduous woodlands in Great Britain and Ireland (Cowling et al. 2003, Vader 1972). Talitroides alluaudi is known outdoors in the Canary Islands, and 7’ topitotum in the Madeira Islands, both species in the Azores (Vader 1972). All other species are generally limited to highly artificial habitats and artificial ecosystems: mostly greenhouses, urban parks and houses (especially cellars). The proportion of introduced isopods can be very high in urban areas. A study in Budapest revealed that 35% of the total species (n = 28) were introduced (Vilisics and Hornung 2009). The major settlements of Hungary were characterised as “hotspot for non-native species” (Hornung et al. 2008). This could certainly be applied to many major cities in other European countries. For the tropical species, especially those recorded only once or twice in Europe, they may not be considered as established (Table 7.1.1.) since their survival is com- pletely dependent on warm greenhouses. Among all alien woodlice, none have spread to more natural habitats. However, the situation is different for intra-European woodlice native to southern or Mediter- ranean Europe. These established aliens can successfully expand by dispersal from very disturbed areas (where they were originally introduced) to more semi-natural habitats in rural-suburban zones (Vilisics and Hornung 2009). With global warming and the large-scale disturbance of biomes in Europe, that trend could increase, especially for the species with large ecological spectra. 88 Pierre-Olivier Cochard et al. / BioRisk 4(1): 81—96 (2010) 7.1.7. Ecological and economic impact of alien terrestrial Crustaceans Alien crustaceans in Europe are not known to carry diseases or to have an impact on native species and natural habitats. Further, they have no economical impact. Based on existing literature, the occurrence of alien woodlice is strictly bound to the urban environment (e.g. greenhouses, botanical and private gardens); alien terrestrial isopods do not yet seem able to survive or to expand to more natural ecosystems. The case of the alien amphipod A. dorrieni is quite different. Terrestrial amphipods are known to have many effects on the soil and leaf litter (Friend and Richardson 1986). Arcitalitrus dorrieni has invaded deciduous and coniferous woodlands in west- ern parts of Great Britain. In Ireland, a study showed that 24.7% of annual litter fall in a coniferous woodland was ingested by this species. It is suggested that “this introduced species plays a more important role than native macrofaunal species in nutrient turno- ver in this particular woodland habitat” (O’Hanlon and Bolger 1999). It is possible that other, as yet undetected, ecological impacts are likely. Terrestrial crustaceans can represent a large percentage of biomass and abundance in the soil macrofauna (Gongalsky et al. 2005). Thus any successful invasion by a ter- restrial alien crustacean could induce some disturbance if it established in relatively natural habitats. For example, in a forested area of Florida, a study on the introduced European woodlouse A. vulgare showed that this species’ activity “had a strong effect on the chemistry of the mineral layer” (Frouz et al. 2008) and concluded that in some cases it may significantly alter soil conditions”. Woodlice classified as aliens of Europe are usually associated with synanthropic habitats and often gain dominance in urban environments (e.g. urban parks, villages, private gardens). The successful colonisation of human- influenced biotopes may lead to the uniformity of local Isopod assemblages. With the decrease of native species in the urban isopod fauna, an ongoing process of biotic homogenisation is prevalent in cities across Europe (Szlavecz et al. 2008, Vilisics and Hornung 2009). Acknowledgements The authors would like to thank Matty Berg (Associate Professor, Vrije Universiteit Amsterdam, the Netherlands); Samuel Danflous (Entomologist, France); Elisabeth Hornung (Associate professor, Szent Istvan University, Hungary); Spyros Sfenthour- akis (Associate professor, University of Patras, Greece); Stefano Taiti (Dr, Istituto per lo Studio degli Ecosistemi, Italy); Ivan H. Tuf (PhD, Palacky University, Czech Repub- lic); and Wim Vader (Professor, Troms@ Museum, Norway). 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