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''Ommatoiulus moreleti'' (spelt ''moreletii'' in older publications), commonly known as the Portuguese millipede, is a herbivorous
millipede Millipedes are a group of arthropods that are characterised by having two pairs of jointed legs on most body segments; they are known scientifically as the class Diplopoda, the name derived from this feature. Each double-legged segment is a resu ...
native to the western
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
where it shares its range with other ''Ommatoiulus'' species. From here, it has spread by international commerce to a number of new localities. This species was accidentally introduced into
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
without its natural enemies and has since become an invasive pest. A number of methods have been developed to manage this millipede.


Distribution

''O. moreleti'' is indigenous to the western Iberian Peninsula. It has spread to a number of Atlantic islands (
Macaronesia Macaronesia (Portuguese: ''Macaronésia,'' Spanish: ''Macaronesia'') is a collection of four volcanic archipelagos in the North Atlantic, off the coasts of Africa and Europe. Each archipelago is made up of a number of List of islands in the At ...
and
Bermuda ) , anthem = "God Save the King" , song_type = National song , song = " Hail to Bermuda" , image_map = , map_caption = , image_map2 = , mapsize2 = , map_caption2 = , subdivision_type = Sovereign state , subdivision_name = , e ...
), South Africa and Australia. This distribution appears to be related to 20th century shipping routes. Spread of ''O. moreleti'' by international trade may be facilitated by its ability to survive long sea voyages in a quiescent state. This species survives long, dry summers in a quiescent state in its natural habitat.Bailey, P.T and Kovaliski, J. 1983. Summer quiescent behaviour of the millipede ''Ommatoiulus moreletii'' (Diplopoda: Julidae). Journal of Zoology London 231: 523-532 After its initial introduction to South Australia in about 1953 (perhaps originating from ships’ ballast) the species is continuing to spread through southern Australia. It is possible that there has been more than one introduction to Australia. Since being introduced to
Port Lincoln Port Lincoln is a town on the Lower Eyre Peninsula in the Australian state of South Australia. It is situated on the shore of Boston Bay, which opens eastward into Spencer Gulf. It is the largest city in the West Coast region, and is located a ...
,
South Australia South Australia (commonly abbreviated as SA) is a state in the southern central part of Australia. It covers some of the most arid parts of the country. With a total land area of , it is the fourth-largest of Australia's states and territories ...
in 1953, the millipede has spread to other parts of
South Australia South Australia (commonly abbreviated as SA) is a state in the southern central part of Australia. It covers some of the most arid parts of the country. With a total land area of , it is the fourth-largest of Australia's states and territories ...
,
Victoria Victoria most commonly refers to: * Victoria (Australia), a state of the Commonwealth of Australia * Victoria, British Columbia, provincial capital of British Columbia, Canada * Victoria (mythology), Roman goddess of Victory * Victoria, Seychelle ...
,
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
,
Australian Capital Territory The Australian Capital Territory (commonly abbreviated as ACT), known as the Federal Capital Territory (FCT) until 1938, is a landlocked federal territory of Australia containing the national capital Canberra and some surrounding townships. ...
, southern
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
and
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
around
Perth Perth is the capital and largest city of the Australian state of Western Australia. It is the fourth most populous city in Australia and Oceania, with a population of 2.1 million (80% of the state) living in Greater Perth in 2020. Perth is ...
.


Life cycle

Reproductive females mature their eggs during late summer-early autumn and may be seen mating during the autumnal activity period after which the female lays 60-80 eggs in a chamber 1–2 cm deep in the soil. The egg hatches to a pupoid stage, then develops by a series of moults up to 16 stages over 3 years. Males can be differentiated by the 8th and 12th stages but most are mature by the 10th or 11th stages. Females probably mature at similar stages. One-year-old immatures (stages 7–9) are light brown with a darker medial stripe. After 2 years the 10-11 stage millipedes have turned black. Adult Portuguese millipedes are smooth, long and coloured from grey to black. Millipedes older than 1 year moult only in spring and summer. Adult males are periodomorphic, alternating between a sexual and a non-sexual form. In their sexual form, they have gonopods (mating legs) in the seventh body segment, which they lose when they moult in spring. They remain in the non-sexual "eunuch" form until their late summer moult.


Activity

The main period of ''O. moreleti'' activity follows the breaking of summer drought by autumnal rains and cooling temperatures. During autumn and early winter, millipedes are active on the surface and may be observed mating. A temperature range 17-21 °C and humidity of around 95% favour activity. These conditions characteristically occur at night. Under similar conditions, individual females on laboratory treadmills move up to 50 metres per day. During winter, there is little surface activity but during spring there is an increase in surface activity. With increasing summer temperatures, ''O. moreleti'' aggregates in humid habitats. At temperatures around 27 °C, millipedes become quiescent.


Habitat

In the southern Iberian Peninsula, a number of species of the genus ''
Ommatoiulus ''Ommatoiulus'' is a genus of millipedes in the family Julidae. The taxonomy of the genus has had a complicated history. As it stands now, there are approximately 60 described species, but this is likely to change. At least 10 new species were d ...
'' share a similar range but divide their habitat on type of litter. This close species packing suggests habitat partitioning may operate to limit numbers of ''O. moreleti'' in relation to other ''Ommatoiulus'' species. ''O. moreleti ''prefers tree litter, particularly ''
Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...
'' spp. (densities stage 7 or older ''O. moreleti'' of about 5 per m²) and ''
Pinus A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
'' spp. whereas ''O. moreleti'' is replaced by other species in shrub litter or grasslandsBailey, P.T and Mendonça 1990. The distribution of the millipede ''Ommatoiulus moreletii'' (Diplopoda: Julidae) in relation to other ''Ommatoiulus'' species in the south-west Iberian Peninsula. Journal of Zoology London 221: 99-101 By contrast, in southern Australia, grassland densities (stage 7 or older) of ''O''. ''moreleti'' of over 40 per m². As an invading species in the southern Australian
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
community, ''O. moreleti'' does not appear to have negatively affected native millipedes sharing a similar range, and seems to have occupied vacant niches.


Food

Gut contents of mature ''O. moreleti'' collected in Portugal were predominantly fragments of ''Quercus'' and ''Pinus'' litter. However, guts also contained significant amounts of fresh
mosses Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and horn ...
and
liverworts The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ge ...
. ''O. moreleti'' can be raised in culture from egg to reproductive stage by feeding solely on fresh mosses.


Natural enemies

Natural enemies in Portugal include the
European hedgehog The European hedgehog (''Erinaceus europaeus''), also known as the West European hedgehog or common hedgehog, is a hedgehog species native to Europe from Iberia and Italy northwards into Scandinavia and westwards into the British Isles.Harris, S. ...
(''Erinaceus europaeus'') and the beetle ''
Ocypus olens The devil's coach-horse beetle (''Ocypus olens'') is a species of beetle belonging to the large family of the rove beetles (Staphylinidae). It was originally included in the genus '' Staphylinus'' in 1764, and some authors and biologists still us ...
'', both of which are generalist predators. A number of parasites have also been identified. None of the natural enemies found in Portugal occur in Australia. In many parts of South Australia, densities of ''O. moreleti'' have declined markedly from a peak period during the 1970s to relatively low densities during mid 1980 to the present. This decline has been associated with parasitism by the
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
''Rhabditis necromena'' that appears to have spread from native millipede populations. This nematode has been actively spread throughout ''O. moreleti'' populations in South Australia by government and private operators. Infection can be detected by crushing a millipede in saline in a petri dish and examining under a X20 dissecting microscope.


Human interactions

In urban areas of southern Australia, ''O. moreleti'' enter dwellings during their autumn and spring activity periods. As a defense mechanism, the millipede secretes a pungent yellowish fluid containing
quinones The quinones are a class of organic compounds that are formally "derived from aromatic compounds uch as benzene or naphthalene">benzene.html" ;"title="uch as benzene">uch as benzene or naphthalene] by conversion of an even number of –CH= group ...
. This stains clothes permanently and irritates eyes. Due to this defence it is best for people to sweep them up rather than crushing them. In South Australia during the 1970s when ''O. moreleti'' were dense in the immediate house surrounds, householders were sweeping volumes of up to several litres from their houses each morning. ''O. moreleti'', alone of the millipede species in this environment, are attracted to low intensity light, of the type emitted from houses at night. Householders in South Australia have used chemical or physical barriers to prevent millipedes from entering houses. Recently, population suppression by spreading the nematode ''Rhabditis necromena'' appears to have effectively reduced millipede populations below worry thresholds over large areas of urban and semi-rural South Australia. In southern Australia ''O. moreleti'' is reported as damaging seedlings of brassica and cereal crops and soft, ripe fruits such as strawberries. Millipedes sheltering in bunches of wine grapes at harvest (autumn) may taint wine. Masses of ''O. moreleti'' on railway lines during their activity periods are reported as causing slippage of locomotives in South Australia, Victoria and Western Australia, causing train cancellations due to the disturbance of signalling equipment. Crushed millipedes may have caused a 2013 minor train crash in
Clarkson, Western Australia Clarkson is an outer northern suburb of Perth, Western Australia, located 34 kilometres north of Perth's central business district in the City of Wanneroo. While mostly a residential area, it also functions as a town centre, serving many surro ...
. Millipedes on the tracks appear to have affected the train's deceleration."Millipedes suspected in Clarkson train crash". Railway Digest (Aust) November 2013 p. 24 In Adelaide, suburban trains are fitted with brushes before the front wheels to clear the line of millipedes.


References

{{Taxonbar, from=Q7090291 Julida Millipedes of Europe Fauna of Portugal Endemic insects of the Iberian Peninsula Animals described in 1860