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Myriapods (Myriapoda). Chapter 7.2 97
Myriapods (Myriapoda)
Chapter 7.2
Pavel Stoev
1
, Marzio Zapparoli
2
, Sergei Golovatch
3
, Henrik Engho
4
,
Nesrine Akkari
5
, Anthony Barber
6
1 National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 So a, Bulgaria 2 Università degli
Studi della Tuscia, Dipartimento di Protezione delle Piante, via S. Camillo de Lellis s.n.c., I-01100 Viterbo,
Italy 3 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospekt 33,
Moscow 119071 Russia 4 Natural History Museum of Denmark (Zoological Museum), University of Copen-
hagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark 5 Research Unit of Biodiversity and Biology
of Populations, Institut Supérieur des Sciences Biologiques Appliquées de Tunis, 9 Avenue Dr. Zouheir Essa , La
Rabta,1007 Tunis, Tunisia 6 Rathgar, Exeter Road, Ivybridge, Devon, PL21 0BD, UK
Corresponding author: Pavel Stoev (pavel.e.stoev@gmail.com)
Academic editor: Alain Roques |Received19 January 2010 |Accepted 21 May 2010 |Published 6 July2010
Citation: Stoev P et al. (2010) Myriapods (Myriapoda). Chapter 7.2. In: Roques A et al. (Eds) Alien terrestrial arthropods
of Europe. BioRisk 4(1): 97–130. doi: 10.3897/biorisk.4.51
Abstract
Alien myriapods in Europe have never been subject to a comprehensive review. Currently, 40 species
belonging to 23 families and 11 orders can be regarded as alien to Europe, which accounts approximately
for about 1.8% of all species known on the continent. Millipedes (Class Diplopoda) are represented by
20 alien species, followed by centipedes (Class Chilopoda) with 16, symphylans with 3 and pauropods
with only 1. In addition there are numerous cases of continental species introduced to the Atlantic and
Mediterranean islands or others of southern origin transported and established in North European cities.
e earliest record of an alien myriapod dates back to 1836, although the introduction of some species
into Europe could have begun already in historical times with an increase in trade between ancient Greeks
and Romans with cities in the Near East and North Africa. In post-medieval times this process should
have intensi ed with the trade between Europe and some tropical countries, especially after the discoveries
of the Americas and Australia. e largest number of alien myriapods (25, excl. intercepted) has been re-
corded from Great Britain, followed by Germany with 12, France with 11 and Denmark with 10 species.
In general, northern and economically more developed countries with high levels of imports and numer-
ous busy sea ports are richer in alien species. e various alien myriapods have di erent origins, but most
of them show tropical or subtropical links (28 species, 70%). Eight of them (20%) are widespread in the
Tropical and Subtropical belts, eleven (circa 28%) are of Asian origin, seven show links with South and
Central America, and one each originates from North America, North Africa, Australasia, and islands in
BioRisk 4(1): 97–130 (2010)
doi: 10.3897/biorisk.4.51
www.pensoftonline.net/biorisk
Copyright P. Stoev 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.
RESEARCH ARTICLE
BioRisk
A peer-reviewed open-access journal
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
98
the Indian Ocean. Ten myriapods are of unknown origin (cryptogenic). Only 12 species (ca. 30%) of all
have established in the wild in Europe. At the present time alien myriapods do not cause serious threats to
the European economy and there is insu cient data on their impact on native fauna and ora.
Keywords
Diplopoda, Chilopoda, Symphyla, Pauropoda, Europe, alien, invasions, intercepted species, biogeograph-
ical patterns
7.2.1. Introduction
Myriapods are terrestrial wingless arthropods with elongated bodies composed of more
or less similar segments, most of which bear one or two pairs of legs. Four classes are
recognised: Pauropoda, Symphyla, Chilopoda and Diplopoda. Approximately 15000
species from nearly 160 families are currently known in the world. e Diplopoda is
by far the most diverse group, comprising roughly 11000 species (Adis and Harvey
2000). A total of 2,245 m yriapod species or subspecies from 15 orders and 70 fami-
lies are currently known in Europe (http://www.faunaeur.org/statistics.php), of which
1,529 are Diplopoda, 481 Chilopoda, 41 Symphyla and 125 Pauropoda. All members
of the class Diplopoda (millipedes) have two pairs of legs per diplosegment for most
segments. Several morphotypes have been recognised, i.e. juloid, polydesmoid, polyx-
enoid, platydesmoid and glomeroid (Kime and Golovatch 2000), of which the former
two are especially rich in species both in Europe and worldwide. Most of the species are
cylindrical or attened dorsally, often with prominent lateral projections, generally me-
dium- to large-sized (up to 8–9 cm in the genera Pachyiulus and Eurygyrus). Some spe-
cies of the order Glomerida, or pill millipedes, are oniscomorph, capable of rolling up
into a tight ball. Members of the order Polyxenida, or “dwarf millipedes”, are minute
in size and with peculiar hairs along the body arranged in groups and tufts like small
pin-cushions or brushes. e number of legs varies between species, often (especially
in juloids) even individually, the record being 375 pairs in the North American sipho-
nophoridan species Illacme plenipes Cook & Loomis, 1928 (Marek and Bond 2006).
Species of the class Chilopoda (centipedes) have an elongated attened trunk and
bear one pair of legs per segment, with a total number ranging between 15 and 191 pairs.
Centipede body length varies from a few millimeters in some species of genus Lithobius
(Monotarsobius) to approximately 30 cm in the Neotropical species Scolopendra gigantea
(Minelli and Golovatch 2001). All centipedes have a pair of poison claws, or forcipules,
which represent modi ed rst body appendages. About 3,500 valid species and subspe-
cies from 5 orders and 22 families are known in the world (Minelli 2006, Edgecombe
and Giribet 2007). e other two myriapod classes – Symphyla and Pauropoda – consist
of very small species, with body length of 2–8 and 0.5–1.9 mm respectively, both still re-
maining very poorly studied. e number of described symphylans and pauropods in the
world is roughly estimated to about 200 and 700, respectively (Adis and Harvey 2000).
Myriapods (Myriapoda). Chapter 7.2 99
Most millipedes, as well as all pauropods and symphylans, are phytophages, detri-
tivores or saphrophages. A few millipedes can be regarded as omnivores, e.g. Blaniulus
guttulatus (Fabricius, 1798), Uroblaniulus canadensis (Newport, 1844), or even preda-
tors, like Apfelbeckia insculpta (L. Koch, 1867), Callipus foetidissimus (Savi, 1819), and
Abacion magnum (Loomis, 1843), which have been observed feeding on earthworms,
ies and spiders (Ho man and Payne 1969, Golovatch 2009). Other species feed on
their own exuvia or fecal pellets (Minelli and Golovatch 2001).
Centipedes are mostly predatory, feeding on di erent available prey items in the
soil (earthworms, enchytraeids, snails, slugs, small insects – both larvae and adults –
and other arthropods). More details on the biology and ecology of millipedes, centi-
pedes and the two other, smaller myriapod classes can be found in Hopkin and Read
(1992), Lewis (1981), and Verhoe (1933, 1934).
Little information is as yet available on the non-indigenous myriapods in Europe
(DAISIE 2009, Roques et al. 2009). e most recent overview of alien organisms
in Europe (see DAISIE 2009, p. 225) lists two centipedes (Lamyctes emarginatus,
Lamyctes caeculus) and three millipedes ( Oxidus gracilis, Eurygyrus ochraceus, Sechel-
lobolus dictyonotus = Paraspirobolus lucifugus) as alien to Europe. Some papers have
been, however, published on the ecology, morphology and post-embryonic develop-
ment of several alien centipedes (Andersson 1984, 2006, Bocher and Engho 1984,
1975a, Negrea 1989) and millipedes (Engho 1975b, 1978, 1987, Golovatch et al.
2000, et al. 2002). Lists of alien species have been published for a few countries
only, such as Germany (Kinzelbach et al. 2001), Austria (Gruber 2002, Gruber and
Christian 2002), the Czech Republic (Šefrová and Laštůvka 2005), Switzerland
(Wittenberg 2005), Italy (Zapparoli and Minelli 2005) and Great Britain (Barber
2009a, b). Increasing attention has been paid in the last decades to species which
have accidentally arrived in Europe (see Barber 2009a, BBC News 2005, Christian
and Szeptycki 2004, Gregory and Jones 1999, Lewis 1988, Lewis and Rundle 1988
for centipedes and Andersson and Engho 2007, Engho 2008a and Read 2008
for millipedes).
7.2.2. Taxonomy of the myriapod species alien to Europe
Altogether, 40 species belonging to 23 families and 11 orders can be regarded as
alien to Europe (Table 7.2.1). is accounts approximately for about 1.8% of all
myriapods known on the continent. Millipedes are represented by 20 alien species,
followed by centipedes with 16, symphylans with 3 and pauropods with only 1. e
relative proportion of alien species is highest in Symphyla (7.3%) and Chilopoda
(3.3%), and the lowest in Pauropoda (0.8%) and Diplopoda (1.3%). e centipede
family Henicopidae is the richest in alien species (5 species), followed by Scutigerel-
lidae, Mecistocephalidae, Scolopendridae, Paradoxosomatidae and Pyrgodesmidae,
each with three species. e remaining 17 families are represented by only one or
two species each (Figure 7.2.1).
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
100
Figure 7.2.1. Relative importance of each family in the alien (right side) and native (left side) myriapod
fauna in Europe. Number near the barindicates the number of species. Families are listed in a decreas-
ingorder based on the number of alien or, in alternative, native species.
Striking is the absence of alien species in Europe of the species-rich order Spiro-
streptida since spirostreptidans, for instance Hypocambala anguina (Attems, 1900) and
Glyphiulus granulatus Gervais, 1847, are quite widespread in the tropical countries and
show a clear tendency to anthropochorism (Jeekel 1963, Shelley 1998).
Several myriapods have been intercepted at their arrival in Europe from consign-
ments from other countries but have never managed to establish themselves. Barber
(2009a) provided a list of centipede species captured and registered by the Central Sci-
ence Laboratory (now FERA) in the UK when imported with exotic plants, fruits and
luggage (Table 7.2.2). Two of them, Lithobius for catus and L. peregrinus, are European
natives which have long been introduced to Australia and New Zealand, so their inter-
ception in Great Britain is a clear case of re-introduction.
A similar list for intercepted millipedes examined by the Central Science Labora-
tories between 1975 and 2006 (S. Reid pers
. comm.) is more substantial with some 85
entries over this period of time (Table 7.2.2). Of these 36 were records of Oxidus graci-
lis from a wide range of di erent parts of the world (W & S. Europe, Canary Islands,
Israel, N., C. and S. America, Australia, China, Japan, Malaysia, Singapore, India,
Nepal, N., W. and S. Africa and Madagascar). Other types found included members
of the Polydesmida (dalodesmids, parodoxomatids, polydesmids), Spirosteptida (from
Australia, New Zealand and Africa), Julidae and Blaniulidae. Amongst species from
the latter two families were the NW European Blaniulus guttulatus and Cylindroiulus
Myriapods (Myriapoda). Chapter 7.2 101
londinensis (both from Australia) and Ommatoiulus moreletii (originating in the Iberian
Peninsula, introduced to Australia in 1953 and now a pest species there; in this list
reported from both that country (tree fern) and South Africa (melon fruit)).
Man-aided introductions of species from one part of Europe to another have played
a prominent role. One of the most common synanthropic centipedes in North Europe
is the Mediterranean “house centipede” Scutigera coleoptrata (Linnaeus, 1758). It has
been introduced to a number of North European cities, e.g., Copenhagen, Edinburgh,
Aberdeen, Leiden, etc., where it survives only in buildings. e earliest record in the
British Isles of this species is perhaps that by Gibson-Carmichael (1883) who recorded
it from a paperworks near Aberdeen. It could have been established there already for 25
years and arrived in bundles of rags from South Europe (Barber 2009a); at the present
time it is sporadically reported from inside buildings in various parts of Britain and
seems to be common in houses in St. Peter Port (Guernsey) and St. Helier (Jersey) in
the Channel Islands from where it has also been reported from outdoor sites. Other
cases of south or central European species being introduced to northern countries that
perhaps still survive only in buildings, hothouses, gardens or similar man-made habi-
tats are: Tuoba poseidonis (Verhoe , 1901) in Finland, Dicellophilus carniolensis (C.L.
Koch, 1847), Lithobius lucifugus L. Koch, 1862, Lithobius peregrinus Latzel, 1880, Ha-
plopodoiulus spathifer (Brölemann, 1897) and Cylindroiulus salicivorus Verhoe , 1908
in Great Britain, Cylindroiulus vulnerarius (Berlese, 1888) in Sweden, Pachyiulus varius
(Fabricius, 1781) in Norway, etc. (Barber 1995, Barber and Eason 1986, Barber and
Keay 1988, Bergersen et al. 2006, Lee 2006, Read 2008).
Even within the same geographic area some indigenous species occur at localities
that are not part of their primary distribution area, most probably as a consequence
of accidental anthropogenic introductions. Examples are the records from Italy of
Lithobius infossus Silvestri, 1894 near Padua (Minelli 1991), of L. peregrinus Latzel,
1880 in northeastern and central Italy (Minelli 1991, Zapparoli 1989, Zapparoli
2006), of Pleurolithobius patriarchalis (Berlese, 1894) in the Egadi, Pontine and Cam-
pania islands (Zapparoli and Minelli 1993), and of Scolopendra cingulata near Milan
(Manfredi 1930).
Island invasions by continental species is another phenomenon worth mention-
ing. Eason in a study on the Icelandic fauna, concluded that most centipede and
millipede species probably arrived by human transport (Eason 1970). Examples of
recent introductions to Iceland are Geophilus truncorum Bergsøe & Meinert, 1866,
Polydesmus inconstans Latzel, 1884, and Brachydesmus superus Latzel, 1884, which “…
have only been found on Heimaey, one of the Vestman Islands, which supports a
town and where casual introduction by human transport is likely: they have probably
been introduced quite recently and the two millipedes seem still to be con ned to
the outskirts of the town”. Regarding the other two possibly allochthonous species,
Lithobius for catus (Linnaeus, 1758), and Lithobius erythrocephalus C.L. Koch, 1847,
Eason wrote, “these two species may be con ned to the south owing to the relatively
warm and humid southern climate, but their restricted distribution might also be
explained by their having been introduced by Norse settlers ....”. e rst Norse set-
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
102
tlements on Iceland were established in the ninth century A.D., so this must have
happened after that time.
According to Engho (2008b), of the 21 species of centipedes recorded in Madeira
17 are introduced and 2 are probably introduced. High rates of introduction are also
known for the Azores and Canary Islands (Borges and Engho 2005, Zapparoli and
Oromi 2004) (Table 7.2.3). All symphylans on the Canary Islands have been consid-
ered as possibly introduced. Likewise, only two of 21 millipede species are considered
native on the Azores (Engho and Borges 2005).
e geophilomorph centipede Nyctunguis persimilis Attems, 1932 was originally
described from Turkey and has not been found there since in spite of the active work
of the second author who has published several papers on the Turkish centipede fauna
during the last 20 years. Taking into account that the species was recently found in
synanthropic habitats in the outskirts of Vienna (Christian 1996) and that all other
congeners occur in the Nearctic region, it is very likely that the type locality (the sur-
roundings of Ankara, Turkey) is erroneous and the material was actually mislabeled
(Zapparoli 1999).
Mecistocephalus maxillaris (Gervais, 1837), one of the rst alien centipedes to be
recorded in Europe, is another poorly known species. It was described from the gardens
of the Muséum National d’Histoire Naturelle, Paris, and subsequently recorded from
numerous places around the world. However, most of the records were certainly based
on misidenti cations with the morphologically similar M. guildingii or M. punctifrons
actually being involved (Bonato et al. 2009). According to Bonato et al. (2009), most
of the records in Europe e.g., those from Germany, Great Britain, France (not the type
specimen but material taken subsequently from a greenhouse in the Paris Museum, cf.
Brolemann 1930) and Portugal (Madeira), are referable to M. guildingii, while those
from the Netherlands and Denmark require further clari cation.
e actual taxonomic status and native range of Ghilaroviella cf. valiachmedovi re-
mains uncertain. e same applies to the millipede Chondrodesmus cf. riparius which
shows some di erences from the original description by Carl (1914) and its identity
cannot be clari ed without a comprehensive review of the entire genus (Engho 2008a).
7.2.3. Temporal trends in the introduction of alien myriapod species to
Europe
Introductions of alien myriapods into Europe probably began several centuries ago,
even though a precise arrival date is hard to determine. Only 10 out of 40 species were
recorded for the rst time in Europe in the 19
th
century while most of the records date
from the 20
th
(26 species) and 21
st
centuries (4 records).
Gervais was virtually the rst person to record alien myriapods in Europe (Gervais
1836, 1837). He described the tropical millipede Iulus lucifugus (now Paraspirobolus
lucifugus) and the geophilomorph centipede Mecistocephalus maxillaris from green-
houses of the Paris Museum. e means of arrival of both species remains obscure but
Myriapods (Myriapoda). Chapter 7.2 103
must be linked to the establishment of the greenhouses and the planting of tropical
owers, perhaps already by the end of the 18
th
century. P. lucifugus has been subse-
quently recorded in intervals of around 60–70 years from greenhouses near Hamburg
(Latzel 1895), Hortus Botanicus Amsterdam (Jeekel 1977), a greenhouse in Copenha-
gen (Engho 1975b), and more recently from the Tropical Biome at the Eden project
(Lee 2006). is can hardly be regarded as re ecting the actual course of colonization
but rather the date of investigation and the level of e ort involved in each case.
e only alien millipede that has invaded some natural ecosystems in Europe and
acclimatized is the East Asian species Oxidus gracilis. Perhaps the earliest records of this
species in Europe are those of Tömösváry (1879) from the Margaret Island in Danube,
Hungary, and of Latzel (1884) from greenhouses in Zeist, Utrecht, and Amsterdam
in the Netherlands. Subsequently it was also found in Edinburgh in 1898 and in Kew
Gardens in Great Britain (Evans 1900, Pocock 1902). In Finland the species was rst
recorded in 1900, but since the sample already contained several specimens the species
must have arrived there at least two years earlier (Palmén 1949). e mechanism of
dispersal of the species within Europe is certainly related to the trading and growing
of tropical plants in the greenhouses as in some places this process must have hap-
pened more than once. According to Palmén (1949), the population of O. gracilis in
the greenhouses of Hanko, South Finland went extinct during the period 1939–1943
when they were not kept warm. In 1946 a single female was found in a greenhouse
with plants imported from Belgium, in 1947 its numbers increased considerably and
the next year it was already very abundant in the whole greenhouse complex.
Golovatch (2008) suggested that the intense trade ties between the ancient town
of Khersonesos in the Crimea and the town of Pergam (= Bergama), a major centre of
red ceramics production of the time in present-day Turkey, as possible pathways for the
introduction of Eurygyrus ochraceus in the Ukraine. He also pointed out that the Bul-
garian population near Varna may owe its origin to the very active commerce in Ro-
man times between Bergama and the colonies in Moesia (= currently northern Bulgaria
and southern Romania), including Odessos (= Varna). e area and trade connections
were already quite developed by the mid-4
th
century B.C. or even earlier, under ancient
Greeks, so this introduction must have happened around that time.
Members of the genus Lamyctes are represented in Europe only by parthenogenetic
populations. Males of L. emarginatus are known only from Macaronesia, New Zealand,
Tasmania and Hawaii (see also Attems (1935) and Zapparoli (2002) for the record of a
single male from Greece), while males of L. coeculus are only known from a greenhouse
in Italy and from Cuba (Engho 1975a). Taking into account that the entire family
Henicopidae is predominantly distributed in the Southern Hemisphere, and presum-
ing that the regions where males are being found are the native areas of the species, L.
emarginatus could have been introduced to Europe from one of the above regions, most
likely from Australia or New Zealand. e earliest con rmed record is from Denmark
in 1868 (see Meinert 1868). Lamyctes coeculus was rst found in a greenhouse in Italy
at the end of 19
th
century (Brölemann 1889), but its presence in the area would have
been older. It has been recently found in Great Britain (Barber 2009b).
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
104
e earliest records of Cylindroiulus truncorum in Europe date from the 1920’s
and, according to Schubart (1925), the Central European populations are probably of
relatively recent origin. In Finland it was rst reported in 1945 and in the following
three years its numbers increased considerably. It is completely lacking in older collec-
tions (Palmén 1949).
One of the recent introductions is the large Neotropical millipede Chondrodesmus
cf. riparius which was rst recorded in 2000 in a owerpot in the telephone o ce of
Umeå University, northern Sweden. It was found again elsewhere in Sweden in 2006
and, later, in January 2007, it was also recorded in a owerpot with a palm (Phoenix
robbelini) in an o ce in Copenhagen and in a owerpot in Bonn (Engho 2008a).
ere are further records of the species from owerpots in Germany and also a recent
one in Norway (Göran Andersson in litt.), so it seems that the species is dispersing well
with palm pots.
e study of the invertebrate fauna of Kew Gardens, Great Britain began al-
ready at the beginning of 20
th
century with papers by Pocock (1902, 1906) and
continues today (Blower and Rundle 1980, 1986, Read 2008). Some of the spe-
cies recorded by Pocock such as Scolopendra morsitans, Trigoniulus corallinus and
Asiomorpha coarctata have not been re-found since then and most likely could not
become established in Kew Gardens. At the same time Paraspirobolus lucifugus, Am-
phitomeus attemsi, Cylindrodesmus hirsutus, Rhinotus purpureus and Pseudospirobolel-
lus avernus, species not previously known from Britain have been recorded recently
in the Tropical Biome at the Eden project in Cornwall (Read 2008, Barber 2009b,
Barber et al. 2010).
7.2.4. Biogeographic patterns of the myriapod species alien to Europe
Records of exotic species are not evenly distributed in Europe but this is mainly due
to the di erent levels of investigation of this area. e highest number of species (25)
has been recorded from Great Britain, followed by Germany with 12, France with
11 and Denmark with 10 alien myriapods (Figure 7.2.2). In general, northern and
economically more developed countries with high levels of imports and numerous
busy sea ports are richer in alien species. ese countries also, in general, have poorer
native faunas meaning that a small number of aliens can constitute a large percent-
age of the fauna. Several species are hitherto known in Europe from a single country
only, e.g. Prosopodesmus panporus, Pseudospirobolellus avernus, Tygarrup javanicus and
Cryptops doriae, which implies recent introductions or poor dispersal abilities. Oth-
ers, such as Eurygyrus ochraceus, Paraspirobolus lucifugus and Lamyctes coeculus, have
a larger but yet fairly restricted distribution limited to two or more countries. e
most widespread species are the parthenogenetic centipede Lamyctes emarginatus,
whose range in Europe spreads from the Urals to Iceland [outdoor species], and
the bisexual millipede Oxidus gracilis, reported from 33 countries, including several
Mediterranean islands.
Myriapods (Myriapoda). Chapter 7.2 105
e various alien myriapods have di erent origins, but most of them show tropi-
cal or subtropical links (28 species, 70%). Eight of them (20%) are widespread in the
Tropical and Subtropical belts, very often introduced by human agency to islands and
synanthropic areas on continents. eir native range cannot so far be determined with
certainty (Figure 7.2.3). Eleven (circa 28%) alien myriapods are of Asian origin, the
majority (10 species) having their native range in East or Southeast Asia, and only one
from West Asia, namely Anatolia. Cylindroiulus truncorum is perhaps the only North
African myriapod introduced to Europe just as Brachyiulus pusillus (Leach, 1814) so
far is the only European julid introduced to North Africa (Akkari et al. 2009). e
only species that seems to be an Australasian native (Australia and New Zealand) is
Lamyctes emarginatus. Among henicopids, Rhodobius lagoi and Ghilaroviella cf. valia-
chmedovi are of particular interest being members of monotypic genera and the only
representatives in Europe of the subfamily Anopsobiidae which comprises chie y spe-
cies with Gondwanan distribution patterns. Besides Rhodobius, four other monotypic
genera represent the subfamily in the Northern Hemisphere, occurring in Vietnam,
Japan, Kazakhstan, and Tajikistan (Edgecombe 2003, Farzalieva et al. 2004). Of Cen-
tral or South American origin are seven species (circa 18%), and one each from North
America and islands in Indian Ocean. e sole record of the pantropical geophilo-
morph centipede Orphnaeus brevilabiatus in Europe comes from Bohuslän, a Swedish
province in the northern part of the W coast, where the animal was collected in the
19
th
century (Andersson et al. 2005).
Ten centipedes and millipedes have been considered as cryptogenic (= species of
unknown origin which cannot be ascribed as being native or alien). Some of them such
as the geophilid Arenophilus peregrinus and the schendylid Nyctunguis persimilis, which
have only been reported from the Isles of Scilly, Great Britain and Austria respectively
(Barber 2008, Christian 1996) whereas all the other species of these genera live in
North America, are of likely Nearctic origins. Another suspected introduction of un-
certain origin is Nothogeophilus turki which has hitherto been known only from Scilly
and the Isle of Wight, Great Britain (Lewis et al. 1988) and represents a monotypic
genus. However, we cannot completely exclude the possibility that some cryptogenic
species suspected to be alien are actually native to Europe. Support for this notion we
nd in the scolopendromorph centipede eatops erythrocephalus C.L. Koch, 1847,
which occurs in various natural habitats (including caves) in the Pyrenees and the
western part of the Balkans (with a gap between these geographic areas), while all its
other four congeners occur in North America (Minelli 2006).
Unknown also is the origin of the symphylid Hanseniella oligomacrochaeta described
from a hothouse in the Botanical Garden in Berlin; according to Scheller (2002), all
species in the genus Hanseniella have tropical-subtropical distributions. e haplode-
smid Prosopodesmus panporus is only known from the Royal Botanic Gardens in Kew,
England, while its other described congener, P. jacobsoni Silvestri, 1910, is pantropical
(Golovatch et al. 2009). Likewise, it is uncertain whether Napocodesmus endogeus, a
millipede described solely from females collected in the garden of Cluj University, is
a European native or not. According to Tabacaru et al. (2003), the generic allocation
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
106
of the second species described in the genus, N. orentzae Tabacaru, 1975, hitherto
known from Romania and Moldova, is not certain and since there are no other records
of N. endogeus in nature it might be an introduced species.
7.2.5. Pathways for the introduction of alien myriapod species in Europe
All of the alien myriapods have most probably been accidentally introduced to Europe
with plant material in relation to human activities and trade between Europe and other
continents such as Asia, Australasia and the Americas. is process must have begun
with an increase in trade between ancient Greek and Romans with cities in Asia Minor
and North Africa and should have intensi ed in post-medieval times with the trade be-
tween Europe and some East Asiatic countries (e.g. Japan, China) and the geographic
discoveries of the Americas and, later, of Australia. is process is still going on with
Figure 7.2.2. Colonization of continental European countries and main European islands by myriapod
species alien to Europe. Archipelago: 1 Azores 2 Madeira 3 Canary islands.
Myriapods (Myriapoda). Chapter 7.2 107
the trade of tropical owers and other plants and their cultivation in houses and green-
houses or with the importing of goods from tropical countries. Even large species could
be transported this way, as is the recent case of the discovery of the largest centipede
Scolopendra gigantea, found in 2005 in a house in London, which is thought to have
arrived with a cargo of electrical goods or fruit (BBC News 2005). Pocock (1906) sug-
gested the possible countries whence a variety of alien species found in Kew Gardens
were introduced with their host plants: India (Scolopendra morsitans, Mecistocephalus
guildingii), Sri Lanka (Chondromorpha kelaarti), Barbados (Anadenobolus monilicornis),
Saint Vincent Island (A. vincenti).
e distribution of the alien diplopods in Europe shows that all the species living
here in greenhouses are much more widespread compared to e.g. the restricted outdoor
species Eurygyrus ochraceus. It is also likely that the obligate thelytokous parthenogen-
esis (= sexual reproduction giving rise to females only) shown in continental Europe
by several of the exotic millipedes and at least one of the centipedes has facilitated
their survival during transport and their establishment on the continent. However,
bisexual populations are known from the Azores and the Canary Islands for Lamyctes
emarginatus (Engho 1975a). Species from other centipede orders, such as the mecis-
tocephalid Tygarrup javanicus also presumably reproduce by parthenogenesis since so
far only females have been found in the hothouse at the Eden project, in Great Britain
(Barber 2009b).
e number of exotic diplopods in Europe is far smaller (3–4 times) than
that of European species introduced to other continents. Apparently, this could
mean that the arrival and, especially, becoming resident in Europe is much more
di cult than the converse process. e asymmetry has probably nothing to do
with quarantine controls at European borders. Instead, it may be due to speci c
ecological and biological patterns exhibited by the successful invaders. Many of
the alien millipedes and centipedes which have successfully invaded Europe be-
Figure 7.2.3. Geographic origin of the myriapod species alien to Europe (in percent).
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
108
long to genera moderately rich to rich in species, such as Poratia, Chondrodesmus,
Lamyctes, Cryptops, etc. A pertinent question arises as to why often only one species
succeeds in establishing populations on foreign continents, sometimes becoming
quite widespread to even cosmopolitan, whereas its rather numerous congeners
fail to do so. Speci c adaptive ecological patterns may be an issue, but, as noticed
Figure 7.2.4. Scolopendra gigantea Linnaeus, 1758 [Chilopoda: Scolopendromorpha: Scolopendridae]
caught in 2005 in apartment in London, perhaps arrived with a cargo of electric goods or fruit. Source:
BBC News: http://news.bbc.co.uk/go/em/fr/-/1/hi/england/london/4201634.stm
Figure 7.2.5. Tygarrup javanicus Attems, 1929 [Chilopoda: Geophilomorpha: Mecistocephalidae].
United Kingdom: Eden Project, Cornwall. Credit: Anthony Barber.
Myriapods (Myriapoda). Chapter 7.2 109
Figure 7.2.6. Rhinotus purpureus (Pocock, 1894) [Diplopoda: Polyzoniida: Siphonotidae]. Japan: Minami-
Daito. Credit: Zoltán Korsós.
Figure 7.2.7. Eurygyrus ochraceus C.L. Koch, 1847 [Diplopoda: Callipodida: Schizopetalidae]. Ukraine:
Crimea. Credit: Kiril Makarov.
above, obligate or opportunist parthenogenesis is probably a major trait favoring
dispersal at least because a single founder juvenile or female is su cient to ar-
rive at destination and found a population. It has to be noted that the successful
myriapod invaders tend to be among the smallest species, thus being more easily
transported, better tted to nd a suitable microhabitat, and sometimes requiring
a shorter time and even a smaller number of developmental stages to reach matu-
rity (Golovatch 2009).
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
110
Figure 7.2.8. Chondrodesmus cf. riparius Carl, 1914 [Diplopoda: Polydesmida: Chelodesmidae]. Denmark:
Copenhagen. Credit: Gert Brovad.
Figure 7.2.9. Oxidus gracilis (C.L. Koch, 1847) [Diplopoda: Polydesmida: Paradoxosomatidae]. Italy:
Porto Badino (Borgo Hermada – Terracina). Credit: Massimiliano Di Giovanni.
Myriapods (Myriapoda). Chapter 7.2 111
Another possible pathway of the introduction of exotic myriapods to Europe is
their intentional import as ‘pets’, and their further escape from pet keepers. Large
Scolo pendra spp., as well as some large and colorful millipedes of the orders Spiroboli-
da, Spirostreptida and Sphaerotheriida are quite popular pet animals subjected to trade
in pet shops. Although there are many guides and internet resources available for keep-
ing and caring for exotic species, there is no reliable information about the importance
of the ‘pet’ trade for the introduction of alien myriapods to Europe. However, the
establishment of pet myriapods in the wild is in most cases very unlikely.
7.2.6. The most invaded ecosystems and habitats
Man-made arti cial environments (pastures and cultivated lands, greenhouses, urban
and suburban areas) constitute the main habitat types hosting alien myriapods (Ta-
ble7.2.1). Species of tropical and subtropical origin are likely to be restricted to green-
houses or equivalent arti cially warmed habitats. Some of them, in the summer season
in the southern countries perhaps could survive also outdoors in close proximity to the
hothouses. However, 11 species have been reported from natural habitats in Europe,
where they most likely were able to establish viable populations. So far the alien spe-
cies of symphylans and pauropods are unknown in natural areas, which is not the case
with several species of the other two myriapod classes. e millipede Oxidus gracilis,
which is bisexual everywhere and is naturalized in several areas in Europe and in the
Caucasus, has been found in forests close to suburban and urban areas (Tömösváry
1879), in woodlands of Robinia pseudoacacia in the Kanev Nature Reserve, Ukraine
Figure 7.2.10. Paraspirobolus lucifugus (Gervais, 1836) [Diplopoda: Spirobolida: Spirobolellidae]. Japan:
Okinawa. Credit: Zoltán Korsós.
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
112
(Chornyi and Golovatch 1993) and records from caves also exist (Strasser 1974, Vicen-
te and Engho 1999). On the Canary Islands the species is quite widespread invad-
ing various, mostly dry and warm, habitats (Arndt et al. 2008). According to Palmén
(1949), O. gracilis dies when subjected for 2 hours to a temperature of minus 4°C.
is means that in North Europe the species can survive only in hothouse conditions.
Cylindroiulus truncorum mainly inhabits synanthropic habitats: greenhouses, gardens,
parks, woodpiles, school grounds, cemeteries, spoil heaps, horticultural nurseries
(Kime 2004, Korsós and Engho 1990).
Eurygyrus ochraceus occurs in the Crimea only in a patch of semi-natural xero-
phytic vegetation ca. 1 km long and 100–300 m wide along a watershed. It was re-
ported to be rather common, although not too abundant on the site and is de nitely
an anthropochore (Golovatch 2008).
Lamyctes emarginatus shows remarkable plasticity regarding the surrounding envi-
ronment, although in the British Isles there is preponderance of rural records in com-
parison with (sub)urban ones. In arti cial habitats it has been reported from gardens,
roads, roadside verges, hedges, embankments, crops of Zea mays and Medicago sati-
va, even in human rubbish (Eason 1964, Minelli and Iovane 1987, Barber and Keay
1988). In natural habitats it lives in various woods (deciduous or mixed coniferous/
deciduous) and has also been recorded from open and coastal areas (Barber and Keay
1988, Zerm 1997, Zapparoli 2006). According to Andersson (2006), it predominates
in open and disturbed areas with sparse vegetation. A great many of these localities
Figure 7.2.11. Trigoniulus corallinus (Gervais, 1847) [Diplopoda: Spirobolida: Trigoniulidae]. Taiwan.
Credit: Zoltán Korsós.
Myriapods (Myriapoda). Chapter 7.2 113
are associated with lake shores, river gravels or river banks. L. emarginatus shows clear
preferences for temporarily ooded sites, no matter for how long the inundation lasts.
Its appearance as a pioneer species on mine sites may indicate that the species shows
preference to disturbed habitats (Zerm 1997). In close proximity to water pools the
species abundance can reach 95% of all centipedes (Minoranskii 1977).
Two of the (presumed) alien geophilomorphs, Arenophilus peregrinus and Notho-
geophilus turki, have been recorded in coastal areas, where they occur under stones and
in soil close to rocky sea cli s with sparse vegetation although A. peregrinus has been
found inland in Cornwall in woodland and one of the Isle of Wight records for Notho-
geophilus turki was from an area of demolished buildings with copious rubbish on the
ground although no more than 5 m from the tidal river (A.N. Keay pers. comm.).
Considerable uctuation in the abundance of some alien species have been observed
by Barber (2009b) in the tropical hothouse of the Eden Project. P. lucifugus which was not
found in 2003/4, was rather restricted in its occurrence in 2005, had become abundant
throughout by 2009. Likewise, C. doriae which has been relatively uncommon and limi-
ted in occurrence in 2005 was the dominant species there in 2009. Conversely, T. java-
nicus, which had been abundant in 2005, was di cult to nd in 2009 (Barber 2009b).
7.2.7. Ecological and economic impact
Alien myriapods are unlikely to pose major threats to native biodiversity and ecosys-
tems. e number of species established in the wild being very limited (12 species, ca
30%) for the moment (Table 7.2.1). Diplopods are detrivorous animals, consuming
10–15% of the leaf litter in temperate forest and as thus contribute signi cantly to
soil formation processes through the fragmentation of leaves which stimulates mi-
crobial activity. ey may thus indirectly in uence the uxes of nutrients (Hopkin
and Read 1992). Nevertheless, some alien diplopods could be harmful to cultivated
plants, especially in the arti cial habitats where temperature and humidity conditions
allow species establishment and expansion. Invasive soil invertebrates may also have
an impact on the structure and function of natural ecosystems. ey can change soil
carbon, nitrogen and phosphorus pools and can considerably a ect the distribution
and function of roots and micro-organisms (Arndt and Perner 2008). In addition,
mass occurrences and swarming, which have been observed in several countries in
Europe, may have negative ecological and economic impact although the causes still
remain obscure (Sahli 1996, Voigtländer 2005). An example of a plant-damaging alien
myriapod is Oxidus gracilis, which is regarded as a pest in several European countries.
is species is very common in greenhouses where its density may exceed 2500 ind./
m
2
. It is known for attacking vegetable and fruit crops such as sugar beet, potatoes,
strawberries, cucumbers, orchard fruits, roots of wheat, and owers in outdoor culti-
vated areas. Furthermore, several thousand O. gracilis were once found after rain in a
house in Lenoir City, Tennessee, USA, with most of the city infested during the same
outbreak (Hopkin and Read 1992). As a curiosity, one might also mention the report
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
114
by the classical writer eophrastus, according to whom an army of millipedes once
overran Rhoeteum in the present province of Çanakkale (northwestern Turkey) and
drove its human inhabitants into the sea (Sharples 1994, Engho and Kebapći 2008).
Several plants can withstand the attacks of symphylans but they may cause severe
damage to growing crops both in elds and greenhouses (Scheller 2002). Arndt and
Perner (2008) recently carried out a study on the impact of invasive ground-dwelling
predatory species, including alien centipedes, in the native laurel forest habitat in the
Canary Islands. ey found that centipedes in laurel forests seem to be much more
variable than carnivorous ground beetles since the 14 recorded species include repre-
sentatives of three orders with very di erent characters. ey tentatively recognised
four functional groups of centipedes: a micro-cephalic schendylid type, (ii) a geophilid
type with medium head size and extreme body length, (iii) a scolopendromorph type,
and (iv) a macro-cephalic lithobiomorph type. ese groups suggest patterns of inva-
sion similar to the coleopteran predators: autochthonous and introduced species of the
same size class and group are mutually exclusive (Arndt 2006).
e potential role of tropical giant millipedes and centipedes (Scolopendra spp.)
kept as pets has been little analyzed as a source of health problems in relation to their
defensive uids or their bites which can cause pathological reactions if exposed to skin,
mouth/throat or eyes (Rein 2002).
Acknowledgements
We thank Helen Read (Farnham Common, UK), John Lewis (Taunton, UK), Greg
Edgecombe (London, UK) and Zoltán Korsós (Budapest, Hungary) for their helpful
comments and shared literature as a result of which the manuscript was able to be
signi cantly improved. John Lewis and Göran Andersson shared unpublished infor-
mation on the presence of alien myriapods in UK and Norway, respectively. Darren
Mann (Oxford, UK) provided a copy of Pocock’s report on Kew species and the paper
by Clarke. We are grateful to Zoltán Korsós, Gert Brovad (Copenhagen, Denmark),
Massimiliano Di Giovanni (Roma, Italy) and Massimo Vollaro (Viterbo, Italy) for
providing photographs of some of the alien species.
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Myriapods (Myriapoda). Chapter 7.2 123
Class
Order
Family Species Status Native range 1st record
in Europe
Invaded
countries
Habitat References
Pauropoda
Tetramerocerata
Pauropodidae Allopauropus pseudomillotianus
Remy & Balland, 1958
A Southeast Asia
(India, Sri Lanka)
1958, FR FR, NO J100 Andersson et al.
(2005)
Symphyla
Symphylomorpha
Scutigerellidae Hanseniella caldaria (Hansen, 1903) A Tropical, subtropical
(North America up to
Mexico, South America,
sub-Saharan Africa, Sri
Lanka, Galapagos Islands,
and possibly New Zealand)
1903, DK DK, FR, GB,
MC, NO
J100 Andersson et al.
(2005), Scheller
(2002), Shear and
Peck (1992)
Hanseniella oligomacrochaeta
Scheller, 2002
C Unknown. Tropical,
subtropical?
2000, DE DE J100 Scheller (2002)
Hanseniella orientalis (Hansen, 1903) A Tropical, subtropical (South
and southeastern Asia,
Central and South America,
islands in the Paci c)
2000, DE DE J100 Scheller (2002)
Chilopoda
Geophilomorpha
Mecisto cephalidae Mecistocephalus guildingii
Newport, 1843
A Amphi-Atlantic (coasts of
Tropical America, African
coast from Gambia to
Liberia, Atlantic islands)
1895, DE DE, FR, GB, PT-
MAD
J100 Bonato et al. (2009),
Pocock (1906)
Mecistocephalus maxillaris
(Gervais, 1837)
C Unknown, tropical? 1837, FR DK, FR, NL J100 Andersson et al.
(2005), Bonato et al.
(2009), Jeekel (1977)
Tygarrup javanicus Atems, 1929 A Southeast Asia (Java,
Vietnam, Cambodia), e
Seychelles
1975, GB AT, GB J100 Barber (2009b),
Christian (1996),
Lewis and Rundle
(1988)
Table 7.2.1. List and main characteristics of the myriapod species alien to Europe. Status: A Alien to Europe C cryptogenic species. Country codes abbreviations
refer to ISO 3166 (see Appendix I). Habitat abbreviations refer to EUNIS (see Appendix II). Only selected references are given. Last update 10/03/2010.
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
124
Class
Order
Family Species Status Native range 1st record
in Europe
Invaded
countries
Habitat References
Geophilidae Arenophilus peregrinus Jones, 1989 C Unknown, genus present in
North America
1986, GB GB B3, I2 Barber (2009a),
Gregory and Jones
(1999), Jones (1989)
Nothogeophilus turki
Lewis, Jones & Keay, 1988
C Unknown 1985, GB GB B3 Barber (2009a), Lewis
et al. (1988)
Oryidae Orphnaeus brevilabiatus
(Newport, 1845)
A Tropical, subtropical
(Australia, Central and
South America, Sub-
Saharan Africa, Madagascar,
East Asia, Arabian
Peninsula, Hawaii)
19
th
century,
SE
SE Un-
known,
J100?
Andersson et al.
(2005)
Schendylidae Nyctunguis persimilis Attems, 1932 C Unknown. Genus present
in North America
1996, AT AT I2? Christian (1996),
Christian and
Szeptycki (2004),
Gruber and Christian
(2002)
Chilopoda
Scolopendromorpha
Cryptopidae Cryptops doriae Pocock, 1891 A Southeast Asia,
Papua New Guinea, e
Seychelles
2007, GB GB J100 Barber (2009a), Lewis
(2007)
Scolopen dridae Scolopendra gigantea Linnaeus, 1758 A Central and South America 2005, GB GB J1 BBC News (2005)
Scolopendra morsitans Linnaeus, 1758 A Tropical, subtropical.
North and South America,
Atlantic Ocean Islands,
Europe, Africa, Arabian
Peninsula, Southeast Asia,
Indian Ocean Islands,
Australia, New Zealand,
Paci c Islands
1902, GB GB J100 Akkari et al. (2008),
Pocock (1906)
Scolopendra subspinipes Leach, 1815 A East and South Asia 1902, GB GB J100 Minelli (2006),
Pocock (1906)
Myriapods (Myriapoda). Chapter 7.2 125
Class
Order
Family Species Status Native range 1st record
in Europe
Invaded
countries
Habitat References
Chilopoda
Lithobiomorpha
Henicopidae Ghilaroviella cf. valiachmedovi
Zalesskaja, 1975
A Unknown. G.
valiachmedovi occurs in
Central Asia (Tajikistan)
2004, AT AT I2 Christian and
Szeptycki (2004)
Lamyctes (Metalamyctes) albipes
(Pocock, 1895)
C Southeast Asia (Java),
Sakhalin Island,
Guadeloupe, e Seychelles
1988, ES-
CAN
ES-CAN H3, H5 Eason and Engho
(1992), Hollington
and Edgecombe
(2004)
Lamyctes (Lamyctes) coeculus
(Brölemann, 1889)
A Tropical, subtropical.
Known from Australia,
Central and South
America, Sub-Saharan
Africa, Madagascar
1889, IT DK, ES-CAN, FI,
FR, GB, IT, SE
J100, J Barber (2009a),
Engho (1975a),
Zapparoli and Minelli
(2005)
Lamyctes (Lamyctes) emarginatus
(Newport, 1844)
A Australasia (Australia+ New
Zealand) is the possible
areas of origin. Known
also from North and South
America, Africa, Asia
Minor, Greenland, Iceland,
New Caledonia, islands in
the Paci c
1868, DK AT, BE, BG, CZ,
DE, DK, ES-
CAN, FI, FR, GB,
GL, GR, HU, IT,
LU, NL, NO, PL,
PT, PT-AZO, PT-
MAD, RO, RU,
SE, SK, UA
B1, D,
E, F4,
F9, G1,
G3, J1,
J2, J3,
J4, J5,
J6, I,
I1, I2,
X6, X7,
X23
Barber and Keay
(1988), Bocher and
Engho (1984),
Meinert (1868),
Minelli and Iovane
(1987), Negrea
(1989), Palmén
(1948, 1952),
Zapparoli and Minelli
(2005)
Rhodobius lagoi Silvestri, 1933 C Unknown, possibly
tropical, subtropical.
Subfamily Anapsobiinae
distributed in South
America, South Africa,
Australia, Japan, Vietnam,
Kazakhstan and Tajikistan
1933,
GR-SEG
GR-SEG
(Rhodes)
I ? Silvestri (1933),
Zapparoli (2002)
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
126
Class
Order
Family Species Status Native range 1st record
in Europe
Invaded
countries
Habitat References
Diplopoda
Polyxenida
Polyxenidae Polyxenus fasciculatus Say, 1821 A Nearctic (USA), Bermuda
Islands
1961, PT-
MAD
ES-CAN, PT-
MAD
G Attems (1935),
Condé (1961),
Vicente and Engho
(1999)
Diplopoda
Polyzoniida
Siphonotidae Rhinotus purpureus (Pocock, 1894) A Tropical, subtropical (South
and Central America,
islands in Indian and
Paci c oceans)
1986, GB GB J100 Barber (2010), Read
(2008)
Diplopoda
Callipodida
Schizo petalidae Eurygyrus ochraceus C.L. Koch, 1847 A Asia (Turkey) 1925, BG BG, UA E1, I2 Golovatch (2008),
Stoev (2007),
Verhoe (1926)
Diplopoda
Polydesmida
Chelodes midae Chondrodesmus cf. riparius Carl, 1914 A South America 2000, SE DE, DK, NO, SE J Andersson and
Engho (2007),
Engho (2008a)
Haplo desmidae Cylindrodesmus hirsutus Pocock, 1889 A Tropical, subtropical (South
America, Southeast Asia,
Papua New Guinea, islands
in Indian and Paci c
oceans)
1950-
1985
AT, DE, FR, GB,
HU, SK
J100 Golovatch and Stoev
(2010), Golovatch et
al. (2001), Golovatch
et al. (2009), Read
(2008)
Prosopodesmus panporus Blower &
Rundle, 1980
A Unknown, other species in
the genus pantropical
1975, GB GB J100 Blower and Rundle
(1980), Golovatch
et al. (2009), Read
(2008)
Myriapods (Myriapoda). Chapter 7.2 127
Class
Order
Family Species Status Native range 1st record
in Europe
Invaded
countries
Habitat References
Onisc odesmidae Amphitomeus attemsi (Schubart, 1934) A South America (Venezuela
or Colombia)
1930, DE AT, CH, DE, DK,
GB, HU, NL,
PL, SK
J100 Barber and Eason
(1986), Engho
(1987), Engho
(2009), Golovatch
et al. (2002), Gruber
(2002), Korsós et al.
(2002)
Paradoxos omatidae Asiomorpha coarctata
(De Saussure, 1860)
A Southeast Asia 1906, GB GB J100 Pocock (1906)
Chondromorpha kelaarti
(Humbert, 1865)
A India, Sri Lanka 1902, GB GB J100 Pocock (1906)
Oxidus gracilis (C.L. Koch, 1847) A Asia (East or Southeast) 1879,
HU
AT, BE, BG, BY,
CH, CZ, DE,
DK, ES, ES-BAL,
ES-CAN, FI, FR,
GB, HU, IE, IS,
IT, LT, LU, LV,
MC, MD, MK,
MT, NL, NO, PL,
PT-MAD, PT-
AZO, RO, RU,
SE, SI, SK, UA
J, J100,
G
Blower (1985),
Engho (2009),
Engho et al. (2004),
Evans (1900),
Ho man (1999),
Pocock (1902), Read
(2008), Šefrová and
Laštůvka (2005),
Stoev (2004)
Pyrgodesmidae Cynedesmus formicola (Cook, 1896) C Unknown, genus native of
Central America
1896, ES-
CAN
ES-CAN, HU,
PT-MAD
J100 Attems (1935),
Korsós et al. (2002),
Vicente and Engho
(1999)
Poratia digitata (Porat, 1889) A Tropical and subtropical
(Southern North and
Central America)
1889, SE AT, CH, DE,
DK, FR, GB, NL,
NO, SE
J100 Blower and Rundle
(1986), Golovatch
and Sierwald (2001),
Gruber (2002), Latzel
(1895)
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
128
Class
Order
Family Species Status Native range 1st record
in Europe
Invaded
countries
Habitat References
Poratia obliterata (Kraus, 1960) A Tropical (South and
Central America: Peru,
Colombia, Brazil, Costa
Rica)
late
1990s,
DE
DE, FR, HU J100 Adis et al. (2000),
Golovatch and
Sierwald (2001),
Korsós et al. (2002)
Trichopoly desmidae Napocodesmus endogeus Ceuca, 1974 C Unknown, only female/s
known; the second
tentative congener occurs in
Romania and Moldova
1969, RO RO I2? Ceuca (1974),
Tabacaru et al. (2003)
Diplopoda
Julida
Julidae Cylindroiulus truncorum
(Silvestri, 1896)
A North Africa (Algeria,
Tunisia)
1925, DE AT, BE, CH, DE,
DK, ES-CAN,
FI, FR, GB, HU,
LT, LU, NL, NO,
PL, PT, PT-MAD,
RO, SE, UA
J, J100,
I2
Engho (2009), Kime
(2004), Korsós and
Engho (1990), Read
(2008), Schubart
(1925)
Diplopoda
Spirobolida
Pseudospiro-
bolellidae
Pseudospiro bolellus avernus
(Butler, 1876)
A Tropical (Southeast
Asia, islands in Indian
and Paci c oceans, and
Caribbean Sea)
2009, GB GB J100 Barber et al. (2010),
Engho (2001)
Rhinocricidae Anadenobolus monilicornis
(Porat, 1876)
A Caribbean region 1906, GB GB J100 Ho man (1999),
Pocock (1906)
Anadenobolus vincenti (Pocock, 1894) A Saint Vincent Island, Lesser
Antilles
1900, GB GB J100 Ho man (1999),
Pocock (1906)
Spirobolellidae Paraspirobolus lucifugus
(Gervais, 1836)
A Tropical. Area of origin
most likely e Seychelles
and/or Mauritius
1836, FR DE, DK, GB, NL J100 Engho (1975b),
Jeekel (2001), Latzel
(1895), Lee (2006),
Read (2008)
Trigoniulidae Trigoniulus corallinus (Gervais, 1847) A Southeast Asia 1902, GB GB J100 Pocock (1906),
Shelley and Lehtinen
(1999)
Myriapods (Myriapoda). Chapter 7.2 129
Table 7.2.2 List of myriapod species intercepted in Great Britain (Barber 2009a, Clarke 1938, John
Lewis, pers. comm., Sharon Reid (FERA), pers. comm.)
Species Native Range Found in/ Country of dispatch/ Year of Interception
Class Chilopoda
Order Craterostigmomorpha
Craterostigmus sp. New Zealand
& Tasmania
Dicksonia (Australia or New Zealand, 2008)
Order Geophilomorpha
? Zelanion (= Steneuryton) sp. Australia, New
Zealand, Hawaii
Dicksonia (Australia, 2005)
Order Scolopendromorpha
Scolopendra cingulata
Latreille, 1829
Mediterranean
region
With luggage (Spain, 2003), potatoes (Greece, 1975),
Turkey (2004), Palestine (pre-1992)
Scolopendra dalmatica C.L.
Koch, 1847
Balkan
peninsula
Found in fruit & vegetable warehouse on Isle of Wight
(1983)
Scolopendra subspinipes
subspinipes Leach, 1815
Asia, Africa,
C. & S. America
Trachycarpus wagnerianus (South Korea, 2006),
bananas (Jamaica, 1938)
Order Lithobiomorpha
Lithobius for catus
(Linneaus, 1758)
Europe Dicksonia (Australia, 2004)
Lithobius peregrinus Latzel,
1880
Europe,
Caucasus
Dicksonia (New Zealand, 2004)
Class Diplopoda
Order Polydesmida
Polydesmida gen. spp. Dracaena fragans (Belgium, 1979)
?Gasterogramma plomleyi
Mesibov, 2003
Tasmania Dicksonia (Australia, 2004)
?Mestosoma sp. Bromeliad (Ecuador, 1982)
Akamptogonus novarae
(Humbert & Saussure, 1869)
? Australia Dicksonia (New Zealand, 2004)
Habrodesmus falx Cook,
1896
West Africa Tete leaves (Nigeria, 1981)
Habrodesmus sp. Orchid (Malawi, 1982)
? Oxidus gracilis ?East Asia Zelkova (Netherlands, 1995)
Oxidus gracilis East Asia Aroid (USA,1980), Chaemaerops (Morocco, 2001),
Cryptomeria (Japan, 1979), Dracaena (Belgium, 1979),
Ficus (West Africa, 1979), Hibiscus (Canary Is.), Lirope
(USA, 1999), Orchid (Belize, 1980; Madagascar,
1995; Malaysia,1984; India, 2000), Palm (Canary
Is., 1998), Pentas (Canary Is., 2010), Phoenix
(USA,
1995), Rhododendron (soil, Nepal, 1981), Sanseviera
(USA, 1980), Scindapus (soil, Nepal, 1981), Selaginella
(Singapore, 1999; Brazil, 1995), Serissa (China,
1999, 2004), Trachycarpus (Netherlands, 2008),
Washingtonia (Italy, 2009), Weeping g (USA, 1984),
Yucca (?Netherlands, 1980), Zamia seed (USA, 1982),
Zelkova (China, 1995), unknown (Chile, 1998; South
Africa, 2001)
Pavel Stoev et al. / BioRisk 4(1): 97–130 (2010)
130
Canary Isl. Azores Isl. Madeira Is. Selvages Isl.
Native Introduced Native Introduced Native Introduced Native Introduced
Diplopoda 83 24 2 19 40 18 2 0
Chilopoda 33* 2** 8 3 2 17+2? 0 2
Symphyla 0 6** 3 0 1 2 no
records
no records
Pauropoda 14*** 0 1 0 10 0 no
records
no records
Table 7.2.3. Relative importance of the non-native species in the myriapod fauna of the Macaronesian
islands. e numbers of introduced species correspond to the total non-native species of both exotic and
continental European origin (cf., Arndt et al. 2008, Baéz and Oromí 2004, Borges, 2008a,b, Borges and
Engho 2005, Engho 2008b, Engho and Borges 2005, Zapparoli and Oromi 2004), some numbers
updated according to recent records. * 7 certainly native, 6 probably native, 20 possibly native, ** all prob-
ably introduced; *** all possibly native.
Species Native Range Found in/ Country of dispatch/ Year of Interception
Polydesmidae Dicksonia (Australia, 2005; New Zealand, 2004),
Orchid (Malaysia, 1983), Wild Plant (South Africa,
1983)
Polydesmus sp. Miscanthus (Dominica, 2000), Orchid (Australia, 1985)
Order Spirostreptida
Spirostreptida Cyathea (New Zealand, 2005), Dicksonia (Australia,
2004–2008), Dracaena (Rwanda, 1980)
Spirostreptus sp. Fig (Ivory Coast, 1983)
Plusioglyphiulus sp. Orchids & Rhododendrons (Borneo, 1979)
Order Julida
Blaniulidae Echinodorus (Singapore, 2008), Orchid (Brazil, 2003)
Blaniulus guttulatus
(Fabricus, 1798)
Europe Orchid (Australia, 1985)
Blaniulus sp. Unknown (South Africa, 1999)
Cylindroiulus londinensis
(Leach, 1814)
Europe Phoenix dactylifera (Italy, 2004)
Cylindroiulus sp. Dicksonia (New Zealand, 2004)
Ommatoiulus moreletii
(Lucas, 1860)
Iberian
peninsula
Dicksonia (Australia, 2006), melon fruit (South Africa,
1983)
Ommatoiulus oxypygus
(Brandt, 1841)
Italy Vitis sp. (Italy, 1979)
Ophyiulus targionii Silvestri,
1898
Italy Unknown (New Zealand, 1982)