Geophilomorph centipedes and the littoral habitat - Books and ...

TAR 

Terrestrial Arthropod Reviews 4 (2011) 17–39 brill.nl/tar 

Geophilomorph centipedes and the littoral habitat 

Anthony D. Barber 

Rathgar, Exeter Road, Ivybridge, Devon, PL21 0BD, England 

e-mail: abarber159@btinternet.com 

Received: 1 October 2010; accepted: 27 October 2010 

Summary 

More than 40 species from at least 20 genera in 6 or more families of the Geophilomorpha (Chilopoda) 

are recorded from marine littoral habitats in various parts of the world. Although there is little recent work 

on their physiology it seems that they have the capacity to tolerate the osmotic and respiratory regime 

that is involved and their anatomical adaptations to a burrowing habit and, at least in some cases, their 

behaviour makes them a fairly constant component of sea-shore ecosystems where they sometimes occur 

in surprisingly large numbers. It is suggested that the richness of the food source in these habitats, along 

with other factors such as shelter, microclimate and possibly absence of parasites and/or predators would 

be the main reason why these now terrestrial animals have re-invaded the seashore so many times since 

their fi rst appearance in the Palaeozoic. Th eir tolerance of seawater and occurrence on coasts could lead 

to passive distribution by rafting and the occurrence of isolated populations could result in genetic 

diff erences. 

© Koninklijke Brill NV, Leiden, 2011. 

Keywords 

Geophilomorpha ; Chilopoda ; littoral habitats. 

Introduction 

Arguing from body fl uid data, Little ( 1990 ) considered myriapods and insects to be of 

marine origin; many legged arthropods would seem to have been some of the earliest 

land arthropods with, so has been suggested, a previous long aquatic history (Selden 

and Edwards, 1989 ). Divergence of centipedes and millipedes has been estimated at 

442 ± 50 Ma (Pisani et al, 2004 ). Terrestrial millipede fossils date back to the Devonian 

(Shear and Selden, 2001 ) whilst evidence for tracheal systems in arthropods comes 

from the Mid Silurian (Wilson and Anderson, 2004 ). Th e earliest centipede remains, 

leg segments of Scutigeromorpha, date from the Upper Silurian (Shear and Selden, 

2001 ). Th ese long-legged animals would have probably have been surface running 

rather than litter inhabiting. It is not until the Late Carboniferous that a possible 

© Koninklijke Brill NV, Leiden, 2011 DOI 10.1163/187498311X546986

18 A.D. Barber / Terrestrial Arthropod Reviews 4 (2011) 17–39 

geophilomorph is recognised in the fossil record (Mundel, 1979 ). Th e divergence 

of the Geophilomorpha is minimally dated by a form from the Upper Jurassic, 

Eogeophilus jurassicus (Shear and Edgecombe, 2010 ). Present day geophilomorphs 

comprise about 1,300 valid taxa, grouped into 14 families representing approxima - 

tely 40% of known centipede species (Rosenberg, 2009 ) and are elongate, worm 

like animals characterised by homonomous segmentation with 27 -191 leg-bearing 

segments. 

Th e earliest report of a halophilic centipede is Leach’s description of the geophilomorph 

now known as Strigamia maritima (Leach, 1817 ). Since its recognition 

further littoral species have been reported; Lewis ( 1981 ) listed 13 and we now know 

of more than 40 taxa from all continents except Antarctica and, of these, all but 

two are geophilomorphs (Barber, 2009a ). Corresponding numbers for the other 

myriapod groups are signifi cantly smaller with only about four to six defi nitely halophilic 

millipedes known (out of a total of about 10,000 species described). Other 

terrestrial arthropod groups also have shore dwelling species but (with the exception 

of Acari) they represent but a small proportion of the known types. According to 

Cheng ( 1976 ), of the species of insects known about 3% are aquatic or have aquatic 

larval stages but of these, only a fraction, “perhaps several hundred species” are marine 

or intertidal. 

At present there are no known species of freshwater centipedes although Strigamia 

maritima has been found in a more or less completely fresh water site (Armitage, 

1982 ); there are two or three millipedes with adaptations for fresh water (Golovatch 

and Kime, 2009 ). Hence the interesting question as to why there are relatively 

so many halophilic geophilomorphs (at least 20 genera in six or more diff erent families; 

Table 1 )? 

Table 1. Geophilomorph families and genera containing known or possible littoral species. (Based on 

Barber, 2009a ) 

Family Genera with littoral species Genera with possible littoral 

species 

Mecistocephalidae Mecistocephalus 

Oryidae Orphnaeus 

Himantariidae Gosothrix, Stigmatogaster 

Schendylidae Hydroschendyla, Nyctunguis, 

Pectiniunguis, Schendyla, Schendylops, Bimindyla, Nesonyx, 

Th indyla 

Ballophilidae Ballophilus, Caritohallex Ityphilus 

Lintotaeniidae Strigamia 

Dignathodontidae Henia 

Geophilidae Erithophilus, Geophilus, Lionyx, 

Maoriella, Mixophilus, Pachymerium, 

Polycricus, Tuoba, Tretecthus 

Diphonyx 

Aphilodontidae Aphilodon

A.D. Barber / Terrestrial Arthropod Reviews 4 (2011) 17–39 19 

Why do terrestrial animals invade the littoral habitat? 

Little ( 1990 ) has a telling diagram (Fig.7.3) of the food web of the thysanuran Petrobius 

brevistylis. Although Petrobius is almost confi ned to the maritime zone and feeds on 

maritime plants its main predators are of terrestrial origin, Lithobius sp. (centipede) 

and Zygiella sp. (spider). 

Here we may have a clue as to why so many diff erent groups of terrestrial animals 

might have invaded the littoral zone. Intertidal areas are often highly productive. Larger 

algae and microscopic forms colonise many seashores and estuarine sites are rich in 

nutrients brought down by rivers. Seashores also receive quantities of (mostly) plant 

material (algae, seagrass, etc.), often in large amounts, brought in by the tides and there 

may be a signifi cant input of guano from seabirds in some locations. In various stages 

of decomposition all this material can provide nutrition for a range of herbivores and 

detritivores which in turn are available to carnivores including centipedes. Th ere are 

various examples in the literature of what appear to be energy fl ow from littoral to 

terrestrial habitats in this way. For instance, Polis and Hurd ( 1995 ) report on extraordinarily 

high densities of spiders on small islands in the Gulf of California which 

they attribute to a combination of energy fl ow from marine to terrestrial food webs 

and the absence of larger predators. Polis et al ( 2004 ) discussed the whole issue of 

trans-boundary transfer between habitats, including that of sea to land. In an extreme 

case of this, Catenazzi and Donnelly ( 2007 ) report on the relationship between the 

terrestrial desert and the highly productive marine environment at Paracas Bay, 

Peru. Consumers in this nocturnal intertidal food web included Collembola, 

Th ysanura, Diptera, Coleoptera, Talitridae, geophilomorph Chilopoda ( Th indyla littoralis 

), Solifugi, Araneae, Scorpionidae and Reptilia. Th e centipedes were recorded at 

an average density of 4.26±0.96 m -1 with a maximum of 49 m -1 . Th e extremely high 

densities of Strigamia maritima sometimes seen on North European shores are very 

striking ( Figure 1 ). 

Even if, perhaps, in other locations, there is actually no net energy transfer between 

marine-littoral and terrestrial food webs at the interface, shoreline production and 

washed up material would broaden the resource base and could entice terrestrial animals 

into the littoral zone. 

In addition to food resources, rocky shores provide crevices which provide shelter 

and could trap air as could burrows in mud and sand. Shingle provides sheltered interstices 

and the drift line a possible rich, if more temporary, shelter and food resource. 

Such environments can provide a degree of protection against weather and predators as 

well as a relatively humid environment and the sea itself will provide an ameliorating 

eff ect on climate in the littoral zone. Th e possible absence of parasites due to unfavourable 

conditions for their alternate hosts might also be a factor in favour of entry of 

species into this habitat (Lewis, 1981 ). 

Geophilomorphs have been observed to feed on a variety of annelids and molluscs 

according to Lewis ( 1981 ). He describes recorded food which including leodocid 

worms (fed on by Hydroschendyla maritima (Grube, 1872)) and unspecifi ed crustaceans. 

Barnacles ( Balanus balanoides ), Orchestia gamarella , Sphaeroma sp. top shells and


Figure 1. Strigamia maritima (Leach, 1817 ) male in shingle Seil Island, Scotland. 

other gastropods ( Littorina saxatalis ) and lumbricid worms were all observed at various 

times to be fed on by Strigamia maritima . Th e same author also described how Strigamia 

maritima attacked small Orchestia (or Drosophila if off ered) by tearing the prey to 

pieces. When attacking larger Orchestia , 1cm or more in length (which it only did if 

they were damaged or dying) it made a transverse slit and pushed its head and anterior 

segments inside and the poison claws were seen to be constantly in motion macerating 

tissue whilst the centipede was involved in what seemed to be external digestion and 

suctorial feeding. Group feeding, as observed in this species could be advantageous in 

dealing with prey that would otherwise be invulnerable to them. Alternatively, small 

specimens could enter barnacles otherwise inaccessible to larger animals and bring 

about their opening. Lewis suggests that it is possible that pheromones might be 

involved in this group behaviour. 

Members of other orders of centipedes such as the scutigeromorphs and lithobiomorphs 

with their relatively long legs and rapid movement would be able to move 

from the supralittoral zone into the upper intertidal area at low tide and back again 

before the tide came in and submerged them or shelter, if necessary, in air fi lled crevices 

as the scorpion Serradigitus littoralis (Williams, 1980) of Baja California is thought to 

do (Due and Polis, 1985 ). Maybe some could even adopt a lifestyle similar to that 

described in the wolf spider Pardosa lapidicina Emerson, 1885 which moves back and 

forth with the tides (Morse, 1997 ). However, such an option would not be available 

to the relatively slow moving geophilomorphs which certainly could migrate up


and down the beach on a seasonal basis as described for Strigamia maritima (Lewis, 

1961a) but would not be able to do this twice a day as required by following the tidal 

cycle; in any case, this would be unlikely to be energy effi cient. If therefore, they were 

able, because of their morphology and physiology, to take up more or less permanent 

residence in areas directly exposed to seawater then this would be advantageous in 

exploiting the littoral resource. 

Adaptation of geophilomorphs to the littoral habitat 

A survey of the locations from which seashore geophilomorphs have been recorded 

includes rock crevices, shingle, under rocks or stones on shingle, sand or mud, under 

stones etc. on salt marshes, under seaweed or other plant material (e.g Posidonia , 

Zostera ) or other material and “strand line debris”. Stranded material off ers a rather 

open habitat and so it is not surprising that it is colonised by a diversity of arthropods 

including centipedes whilst crevice dwelling off ers great opportunities for the more or 

less burrowing geophilomorphs with their worm-like bodies and their modes of 

locomotion. 

Animals living here need to be able to tolerate (or avoid by migration or survival in 

air spaces) the eff ect of inundation with its consequential respiratory and osmoregulatory 

implications. Animals will be immersed in seawater twice daily for a shorter or 

longer time depending on their location on the shore although not normally facing 

continuous 24 hour immersion. Th ey will also be in contact with saline substrates 

(algae, sand, stones, etc.) and will be taking in salt ions in their food. In estuarine 

conditions there will also be an issue of varying salinity, generally below that of full 

sea-water. 

Unfortunately most of the work on the physiology and adaptations of littoral 

geophilomorphs is rather old and further studies are clearly desirable. 

Body design 

Manton ( 1965 ) identifi ed a range of features that adapted geophilomorphs to their 

burrowing and crevice dwelling life and, although she generalised from studies on a 

small number of species, her observations do seem to have some relevance. However, it 

must be borne in mind that there is considerable variation as between diff erent species, 

genera and families. For instance, the head is more or less oblong rather than tapered 

in some species (e.g. some Geophilu s spp., Pachymerium ferrugineum ) nor do all show a 

clear ‘transition’ and members of the Mecistocephalidae are unlike other geophilomorphs 

in a number of ways. 

When walking, geophilomorphs proceed relatively slowly although with little lateral 

undulation, feeling their way forward with their antennae (all are eyeless) and with 

the capacity for backward movement if required. Th is allows them to explore cre - 

vices in search of prey or shelter and to retract as necessary and when sheltering their 

fl exible multi-segmented bodies allow them to coil up and occupy a limited volume 

( Figure 2 ).


Figure 2. Hydroschendyla submarina (Grube, 1872) as found coiled up in a rock crevice at Wembury, 

England. 

When burrowing in an earthworm-like manner, the large number of short segments 

each of which can shorten and thicken allows extension and contraction of trunk 

regions. In addition the head is relatively small and often tapered with mouthparts 

which sweep semi fl uid material from their partially externally digested prey into 

the digestive system. Th e anterior third of the trunk (in front of the so called “transition”, 

if present) may be very distinctively tapered and with the narrow oesophagus, 

heart and nerve cord having been described as centrally positioned to minimise deformation 

by changes in the dimensions of the segments. Manton describes the cuticle 

as being thicker in this anterior region and forming an armour of sclerites including 

intecalary tergites and sternites which allow for changes in shape whilst at the same 

time individual sclerites have fl exibility at the margins and stiff ness in the centre so 

that thrusts can be transmitted but the armour remains intact and the plates do not 

crack. In addition the longitudinal muscles are relatively large. An anterior region of 

the body of the head and anterior trunk with relatively stout legs, adapted for strong 

slow movements, allows extension and contraction of the front of the body with headon 

burrowing to make the burrow wide enough for the remainder of the body to 

follow. ( Figure 3 )

Body fl uid regulation 


Figure 3. Stigmatogaster subterranea (Shaw, 1789) to show the body shape of a geophilomorph with a 

distinct transition. Although normally a terrestrial species, this specimen was actually found under a stone 

on estuarine mud, River Lynher, Cornwall, England. 

As a legacy of their original marine origin myriapods have body fl uid concentrations 

comparable to that of crustaceans (Little, 1990 ). For the two marine species studied by 

Binyon and Lewis ( 1963 ), Strigamia maritima has a body fl uid OP of 462 mOsm.kg - 

and a Na + level 274 mM.l -1 and Hydroschendyla submarina 498 mOsm.kg - and 283 

mM.l -1 which are only slightly higher than the terrestrial Stigmatogaster subterranea 

(Shaw, 1789) with 438 mOsm.kg - and 205 mM.l -1 although higher than those of the 

lithobiomorph Lithobius forfi catus at 370 mOsm.kg - and 190 mM.l -1 (Little, 1983 ). 

Th ese levels are about half or slightly less than that of full seawater so that such littoral 

animals will still need to have mechanisms for regulating body fl uid volume e.g. 

removal of excess ions by some mechanism. Alternatively, they will need be able to 

tolerate changes of body fl uid concentration. 

For terrestrial centipedes with their incomplete waterproofi ng / limited spiracular 

closing mechanisms, dry air poses a threat of desiccation (as would seawater). Damp 

environments which they normally tend to inhabit will pose the opposite threat of 

osmotic water uptake which seems to be dealt with by removal of water in the gut and 

retention of ions. It is possible that the coxal glands may, in some way be involved in 

transporting (Little, 1983 ); see also the review of these in Rosenberg ( 2009 ).


Geophilomorph centipedes, both terrestrial and littoral have been shown to be 

more tolerant of immersion than lithobiomorphs (Hennings, 1903 Plateau, 1890 ). 

Schubart ( 1929 ) found that submerged specimens of Pachymerium ferrugineum 

(C.L.Koch,1835) from Germany survived 7-68 days at room temperature (16-18 o C) 

whilst Suomalainen ( 1939 ) reported 24-95 days at 19-27 o C and 68-178 days at 6-12 o C 

for Finnish animals. Binyon and Lewis ( 1963 ) reported on survival times for animals 

immersed in sea-water for Strigamia maritima , Hydroschendyla submarina 

Stigmatogaster subterranea with the latter having much reduced survival times compared 

with the two littoral species. Th e two halophiles were able to maintain their body 

fl uid concentrations more or less constant for 14 hours whilst in Stigmatogaste r this 

rose signifi cantly. Th ere appears to have been little subsequent work on osmoregulation 

in littoral centipedes. 

Gaseous exchange 

Unlike some spiders which are able to trap bubbles of air in their nests and use these 

either as a direct resource or as a physical gill or remain in air pockets trapped in some 

other way e.g. in empty barnacle shells (McQueen and McLay, 1983 ; Roth and Brown, 

1975 ; Rovner, 1987 ) there is little clear evidence of geophilomorphs doing this even 

though Bonnel ( 1929 ) had reported that Mixophilus indicus (Silvestri, 1929 ) trapped a 

bubble of air in a loop in the posterior end of its body. Rajalu ( 1972 ) was unable to 

confi rm this and as Lewis ( 1962 ) pointed out, although newly immersed specimens of 

Strigamia maritima have bubbles of air on their surface these are likely to be rapidly 

dispersed by movement. Nor is there any evidence of centipedes occupying air fi lled 

burrows. However there have been suggestions that air bubbles could be trapped in 

spiracles and might act as physical gills (Laloy, 1904 ; Suomalainen, 1939 ) and there is 

some evidence for this (Lewis, 1960). 

Alternatively either the general body surface or some part of it might allow gaseous 

exchange directly with sea water (Rajalu, 1972 ). Manton ( 1965 ) argued that closure of 

the spiracular atrium when burrowing allowed respiration to continue and that the 

atrial cuticle facilitated respiration under wet soil (and hence, possibly, sea water) 

conditions. 

Tolerance of more or less severe hypoxia in insects (which would be advantageous 

during submergence) is greater than in most vertebrates (Hoback and Stanley, 2001 ; 

Schmitz and Harrison, 2004 ) and the apparent capacity of at least some centipedes to 

build up an oxygen debt which is subsequently “paid off ” is indicated by experiments 

with Mixophilus indicus (Rajalu, 1970 ). A similar phenomenon has been found when 

measuring oxygen consumption in the terrestrial lithobiomorph Lithobius agilis (Ivan 

Kos, pers. comm.) 

Problems of defi ning halophiles 

One of the problems associated with gaining a clear picture of the variety and numbers 

of shoreline animal species is that of defi nition of the terms “littoral” and “sea shore” 

which, in some accounts, can include areas of land close to the beach itself. For instance, 

and


Ivinskis and Rimšaitė ( 2005 ) in a study of the Curonian Spit and Klaipėda-Šventoji 

zone refer to “seashore habitats” in which they included dunes, dry grasslands and sand 

heaths (and recorded 2,000 species of insects). Correspondingly “Littoral Méditerranéan” 

as used by Brölemann ( 1930 ) when describing some geophilomorph species seems to 

refer to that particular region of France rather than to specifi cally littoral habitats as we 

might understand them. 

In this account we will take the term to refer to species found in locations close to 

high water line and below. Including the supralittoral zone would substantially increase 

the number of species as more “terrestrial” forms are found but the line between littoral 

and supra-littoral species is necessarily blurred. Lewis ( 1962 ), when describing the 

ecology, distribution and taxonomy of the centipedes on the shore in the Plymouth 

area, includes four species that we might regard as typically littoral [ Hydroschendyla 

submarina , Schendyla peyerimhoffi (Brölemann and Ribaut, 1911), Strigamia maritima , 

Geophilus seurati (Brölemann, 1924; Figure 4 )], a species which occurs both on and off 

the shore ( Geophilus fl avus ) and one, Stenotaenia linearis (C. L. Koch, 1835), which is 

usually regarded as terrestrial and was found at high water mark. 

In addition to the problem of defi nition, littoral habitats are, by their nature, often 

diffi cult to sample, whether shingle, saltmarsh, sand, mud, mangroves or transitory 

drift lines and in addition to the physical diffi culties in extracting littoral species from 

their substrate some appear to have discontinuous distribution patterns and even 

within a particular location, possibly because of micro-variation in environmental 

conditions, they may show a patchy occurrence. Th ese together with tidal, seasonal or 

Figure 4. Geophilus seurati Brölemann, 1924 (= G.gracilis Meinert, 1898) from under a stone on a salt 

marsh, Gower Peninsula, Wales.


weather induced local migration can sometimes make it surprisingly diffi cult to fi nd 

littoral species in apparently suitable sites. 

Categories of halophiles 

Th ere are clearly and distinctly recognised halophilic species such as the European 

Hydroschendyla submarina which are only found on shores below or around high water 

line and which will be likely to experience submersion at high tide but species found 

on the shore may also include typical “terrestrial” ones that happen also to occur sometimes 

on the shore during chance foraging but are not specifi cally associated with it 

and this could include a very wide range of types. Th ere are also species that do commonly 

occur on the shore and are characteristically found there but also occur inland 

such as Pachymerium ferrugineum widespread across Europe and elsewhere but regularly 

found in beach shingle or other littoral sites e.g. in Finland (Palmén, 1949 ). 

Silvestri ( 1903 ) distinguished these three categories as miriopodi halofi li genuini , 

miriopodi halofi li accidentali and miriopodi halofi li indiff erenti and these remain useful 

concepts. Unfortunately there are many species described which, one suspects, could 

be littoral but lack detailed habitat data. Th ere are others where there are so few records, 

possibly only a single one so that it is impossible to which of Silvestri’s categories the 

apparently littoral species might be allocated. Th ere are also animals which appear to 

be genuine halophiles in one region but have been found inland elsewhere. Examples 

of the latter include Schendyla peyerimhoffi and Schendyla monodi (Brölemann, 1924), 

both apparently exclusively halophilic in NW Europe but which have been recorded 

inland in Iberia. 

Th e range of species and their known geographical distribution 

Littoral geophilomorphs, whether genuini , indiff erent i or possibly accidentali have 

been described from many seashores, sometimes only once but there are areas of the 

world such as much of Asia, sub-Saharan Africa and the Atlantic coast of North 

America which are poor in known species, possibly due to lack of collection and identifi 

cation from suitable sites or lack of suitable habitats. Th e present list ( Tables 1, 2a, 

and 2b ) is based on the literature, mostly as summarised with the relevant references in 

a previous paper by the present author (Barber, 2009a ) where are also listed other possible 

littoral species for which there is inadequate habitat data. In the tables the probable 

status is indicated on the basis of known ecological data (G = genuini , I = indiff erenti , 

R = littoral in one region, inland in another) but it is quite possible that some of the 

myriopodi halofi li genuini may turn out to be indiff erenti or even accidentali . 

It is also possible that not all species names are valid either because of synonymy or 

lack of clarity as to whether the name is being correctly applied. In an example of the 

latter, Chamberlin ( 1909 ) described a new species Pectiniunguis heathi (now known as 

Nyctunguis heathi ) as up to 22mm and from a shell mound near Cypress Point, Montery 

County, California without actually indicating whether this was a shoreline site.


Table 2a. Littoral geophilomorph species and their recorded distribution: Families Mecistocephalidae, 

Schendylidae, Ballophilidae, Linotaeniidae, Dignathodontidae, and Aphilodontidae. Based on data in 

Barber, 2009a. 

FAMILY and Species Distribution Status Note 

MECISTOCEPHALIDAE 

Mecistocephalus manazuensis Shinohara, 1961 Japan ? I 1 

Mecistocephalus satumensis Takano, 1980 

SCHENDYLIDAE 

Japan G 

Hydroschendyla submarina Grube, 1872 Mediterranean, N.Atlantic 

(Scandinavia–N.Africa), 

Bermuda 

G 

Nyctunguis heathi (Chamberlin, 1909 ) California ?G 2 

Pectiniunguis albermarlensis Chamberlin, 1914 Galápagos Islands I 

Pectiniunguis americanus Bollman, 1889 Gulf of California ?I 

Pectiniunguis amphibius Chamberlin, 1923 Gulf of California, Florida, 

Mexico 

G 

Pectiniunguis bollmani Pereira et al, 1999 Venezuela G 

Pectiniunguis halirrhytus Crabil, 1959 Florida, Mexico G 

Pectiniunguis krausi Shear and Peck, 1992 Galápagos Islands I 

Schendyla monodi (Brölemann, 1924) France 

Inland in Spain 

R 3 

Schendyla peyerimhoffi (Brölemann and Great Britain, Ireland, France R 

Ribaut, 1911) 

Inland in Portugal, Algeria 

Schendylops virgingordae Crabill, 1960 Virgin Gorda, Martinique, 

Venezuela 

G 

Th indyla littoralis (Kraus, 1954) 

BALLOPHILIDAE 

Peru G 

Ballophilus riveroi Chamberlin, 1950 Tortola (W.I.) ?I 4 

Caritohallex minirrhopus Crabill, 1960 

LINOTAENIIDAE 

Tortola (W.I.) G 

Strigamia maritima (Leach, 1817 ) Scandinavia – N. France 

Great Britain and Ireland 

G 

Strigamia japonica (Verhoeff , 1935 ) 

DIGNATHODONTIDAE 

Japan G 

Henia bicarinata (Meinert, 1870) 

APHILODONTIDAE 

Mediterranean, Macaronesia, 

Canary Islands 

I 

Aphilodon maritimus (Lawrence, 1963) South Africa ?G 5 

Status: G = genuini , I = indiff erenti , R = littoral in one region, inland in another. 

Notes: 

1. May be synonymous with M. nannocornis Chamberlin, 1920, most records of which are non-coastal 

(Uliana et al., 2007 ). 

2. See comment about this species. 

3. May be synonymous with S. viridis (Verhoeff , 1951). 

4. Male agreeing with this description collected at same time by same method as C. minirrhopus. 

5. Included on basis of name and location but not specifi cally identifi ed as littoral.


Table 2b. Littoral geophilomorph species and their recorded distribution: Family Geophilidae. Based on 

data in Barber, 2009a and named sources. 

Species Distribution Status Note 

Erithophilus neopus Cook, 1889 Florida ?I 1 

Geophilus admarinus Chamberlin, 1952 Alaska G 

Geophilus algarum Brölemann, 1909 France (north and west coasts) G 2 

Geophilus becki Chamberlin, 1951 California G 

Geophilus fl avus (De Geer, 1778) Widespread in Europe, 

Newfoundland, North America 

I 

Geophilus fucorum Brolemann, 1909 Mediterranean G 2 

Geophilus proximus C.L. Koch, 1847 Central and Northern Europe 

Fennoscandia, White Sea 

3 

Geophilus pusillifrater Verhoeff , 1898 Great Britain, Ireland, Brittany 

Inland in Herzegovina 

R 4 

Geophilus seurati Brölemann, 1924 Great Britain, Ireland, Brittany 

G 2, 5 

(= G. gracilis Meinert, 1898) 

Mediterranean (North Africa), USA 

Geophilus terranovae Palmén, 1954 Newfoundland 6 

Lionyx hedgepethi Chamberlin, 1960 California G 

Maoriella ecdema Crabill, 1964 Chatham Island (New Zealand) ?G 7 

Mixophilus indicus (Silvestri, 1929 ) India G 

Pachymerium ferrugineum 

Europe, North Africa, Taiwan, 

I 8 

(C.L.Koch, 1835) 

Hawai’i, Japan, North America, etc. 

Polycricus bredini (Crabill, 1960 ) Tortola (West Indies) ?G 9 

Tretecthus uliginosus (von Porat, 1894) Cameroun G 

Tuoba ashmoleorum (Lewis, 1996 ) Ascension Island ?G 

Tuoba japonicus (Fahlander, 1934) Japan G 

Tuoba kozuensis Takakuwa, 1934 Japan G 

Tuoba laticeps Pocock, 1901 Australia (Western Australia, Tasmania) G 

Tuoba littoralis (Takakuwa, 1934) Japan G 

Tuoba pallida Jones, 1998 Western Australia G 

Tuoba posedonis (Verhoeff , 1901) Mediterranean, Dead Sea, Red Sea G 

Tuoba sudanensis (Lewis, 1963) Sudan G 

Tuoba sydneyensis (Pocock, 1891) Australia, New Caledonia, New Britain, G / I 10 

Solomon Islands, Hawai’i 

Tuoba tiosanus Takakuwa, 1934 Japan, Micronesia ?G 

Tuoba xylophaga (Attems, 1903) New Zealand G 

Status: G = genuini , I = indiff erenti , R = littoral in one region, inland in another. 

Notes: 

1. Included on advice of L. A. Pereira. 

2. See comment about these species. 

3. Littoral in Fennoscandia (Palmén, 1949 ), White Sea (along with P. ferrugineum ; Prziboro, 1994 and 

pers. comm.). 

4. Described originally from inland site in Herzegovina. 

5. Th ere is a record of Geophilus gracilis from New Haven, Connecticut (Harger, 1872 ). 

6. A number of seashore records, mostly or entirely above high water mark (Palmén, 1954 ). 

7. Under stones, high water mark (Crabill, 1964 ). 

8. Pachymerium idium Chamberlin, 1960 was taken in a rock fi ssure kept moist by spray but not covered 

by high tide, California (Chamberlin, 1960 ). 

9. Berlese sifting of beach drift. 

10. Occurs up to 950m in Hawai’i.


Evans ( 1980 ) in an accounts of littoral arthropods from California describes this species 

as occurring “in crevices and on the surfaces of rocks at night during low tide” and 

up to 45 mm long and in a similar record of its occurrence on the shore by Ricketts et 

al ( 1985 ) it is described as “common”. A photograph in the book Living Invertebrates 

(Pearse et al, 1987 ) shows a 50 mm specimen identifi ed as N. heathi . In relation to this, 

Hoff man and Carlton ( 2007 ) commented that “Th e identifi cation of any schendylid 

should be approached with caution however and the identity of Chamberlin’s species 

with intertidal centipedes may bear re-examination”. 

In the case of Geophilus fucorum Brölemann, 1909, Geophilus seurati Brölemann, 

1924 (= Geophilus fucorum seurati Brölemann, 1924, also known as G. gracilis Meinert, 

1898), Geophilus algarum Brölemann, 1909, and G. algarum decipiens Brölemann, 

1930 , there are issues relating to taxonomy / nomenclature and to variability within 

and this group (Lewis, 1962 ) and examination of further specimens from both the 

British and French coasts would be helpful. 

Orphnaeus brevilabiatus (Newport, 1845) (Family Oryidae) is included in a list of 

littoral species by Roth and Brown ( 1976 ) who quote Crabill (pers.comm.) describing 

it as being found in “cocoon-like structures” on twigs fl oating or awash on beaches. 

It is a species widespread in the tropics and if this is the case then this might be a way 

in which it is dispersed and it might be considered an indiff erent halophile. 

Stigmatogaster subterranea (Family Himantariidae) a north-west and central European 

terrestrial and often synanthropic species has sometimes been found on the upper 

shore in company with Strigamia maritima or Geophilus seurati on the western coast of 

Great Britain (personal observations) ( Figure 3 ). Th is is despite the fact that it has a low 

survival time in seawater (Binyon and Lewis, 1963 ). 

Hydroschendyla submarina occurs on the eastern Atlantic coast from Scandinavia to 

North Africa in crevices at or below high water mark and in the Mediterranean (France, 

Greece, Italy and North Africa) from where it is recorded under stranded remains of 

Posidonia and near salt marshes. It is also known from Bermuda in crevices in the upper 

intertidal and low supra-littoral. Lewis ( 1962 , 1981 ) contrasted its behaviour with that 

of Strigamia maritima and also showed that, having eggs impermeable to sea water 

allowed both moulting and egg laying in the littoral zone. 

Strigamia maritima is recorded from the coasts of Scandinavia, Germany, Netherlands, 

Belgium, Britain, Ireland and northern France. Bergesen et al. (2006) recorded it for 

the fi rst time for North Norway (under stones in the supralittoral amongst isopods, 

Porcellio scaber ) whilst Horneland and Meidell ( 1986 ) were able to collect “several 

thousands of specimens” from an island north of Bergen. It is often extremely abundant 

in favourable situations with large numbers under individual stones, a situation 

that seems to be paralleled by Geophilus becki according to Habermann (1982). Habitats 

for S.maritima include shingle, under rocks and stones and in rock crevices. Th e life 

history and ecology of this species was studied by Lewis (1960, 1961 ) who related its 

behaviour and its tolerance of seawater at diff erent stages of its life history to it being a 

mobile species, concentrating in areas that are climatically favourable and have a good 

food supply and to its breeding season and its ability to migrate up and down beaches.


Pachymerium ferrugineum is cited as having coastal areas and particularly seashores 

as a “preferred” habitat at least in northern Europe but certainly not being an exclusive 

shore species and with a wide ecological range, found up to 2,800m asl in the Hoggar 

Mountains in the Sahara (Palmén and Rantala, 1954 ). It is possibly one of the most 

widespread centipedes in the world and is known from Scandinavia, Netherlands, 

Belgium, France, Iberia, Mediterranean region including North Africa, Asia Minor, 

Caucasus, Azores, Canaries, Madeira, Taiwan, Japan, Hawai’i, Juan Fernandez, Mexico, 

USA, Canada, Alaska, etc. Th ese are certainly not all littoral sites and in some places 

it is no doubt present because of accidental human introduction but its physiological 

tolerance certainly suggests the possibility of “natural” expansion of its range e.g. by 

oceanic rafting to isolated islands and coastal locations. Mediterranean records include 

a variety of habitats including the seashore and in wrack. In the Black Sea (Bulgaria) it 

is recorded (as var. insulanum ) from under stones, algae and Zostera . 

In northern and western Europe its distribution is interesting. It is known from 

Norway, Sweden, Finland and Denmark and from the White Sea coast as well as inland 

in NE Europe. In Eastern Fennoscandia records include from under stones, in decaying 

Fucus and other debris on seashores as well as from dry terrestrial sites (Palmén, 

1949 ). Meidell ( 1979 ) drew attention to the fact that this species and Strigamia maritima 

form an “east-west pair of species” with S. maritima along the western coast and 

P. ferrugineum along the eastern coast of southern Norway and a zone of overlap in 

both the north and south. Bergersen et al . ( 2006 ) did not list it for north Norway. Th e 

maps for the two species in the Encyclopedia of the Swedish Fauna and Flora (Andersson 

et al. , 2005 ; pp. 165 and 169) highlight this pattern in the Nordic countries. 

All records from the Netherlands and Belgium appear to be from inland sites 

(Berg et al., 2008 , Lock, 2009 ). It has described as widely distributed in France and 

Corsica and favouring the seashore “without being strictly halobiontic or highly halophilous” 

(Geoff roy and Iorio, 2009) but the only record from the northern coastal area 

seems to be from a seashore dune in Brittany (Iorio, 2005 ). Th e three British records 

are all from beach shingle (Barber, 2009b ). Th is would suggest that in this area of 

Western Europe it is very much on the edge of its European range and that maybe 

competition with each other in some way restricts the range of P. ferrugineum 

and S. maritima . Interestingly, Andersson ( 1983 ) noted a substantial decrease in 

P. ferrugineum in open rocky terrain, on seashores and the Northern Skerries between 

the 1920s and the 1970s in the vicinity of Göteborg (Sweden). No explanation of this 

decline was suggested. 

Characteristics of littoral geophilomorphs 

Examination of the range of littoral Geophilomorpha reveals an extremely diverse 

group of species. For instance, bearing in mind that most measurements are based on 

preserved material, recorded body lengths for littoral forms vary from 10mm ( Caritohallex 

minirrhopus ) to 55mm ( Th indyla littoralis , Henia bicarinata ;) and leg-bearing 

segments from 39 ( Schendyla peyerimhoffi , C. minirrhopus ) to 85 ( H.bicarinata ).


Th e actual number of leg-bearing segments varies slightly as between sexes and within 

most species (apart from mecistocephalids) but remains constant throughout life. Body 

colour varies between pale / semi-transparent through various shades of yellow and 

orangish to reddish-brown with head region often a somewhat darker although colours 

often fade in preserved specimens. Mixophilus indicus was described as “light leather 

coloured” by Silvestri ( 1929 ). 

Crabill ( 1968 ) drew attention to the fact that in the genus Tuoba (Geophilidae) 

which contains a number of littoral species there was a long and spiniform pre-tarsal 

anterior parunguis (spine) on the legs which he regarded as associated with the genus’s 

distinctively littoral preferences and which, he suggested, probably functioned as a 

special hold-fast adaptation to which Lewis ( 1996 ) added the comment that it would 

also be useful during transportation. Pereira et al. ( 1999 ) reported on a similar structure 

in Pectiniunguis (Schendylidae). Other than this, across the various species and families, 

there seem to be few common characteristics that are obviously related to a littoral 

lifestyle and even within a single family there may be marked diff erences between different 

littoral forms. For instance, in the Schendylidae, Hydroschendyla submarina is a 

relatively robust reddish brown form up to 40mm long, Schendyla peyerimhoffi is pale 

yellow and up to 18mm whilst Th indyla littoralis is described as yellowish with head 

orange and up to 55mm long. Closer examination and comparison might, perhaps, 

identify common features shared by many or all littoral forms. 

Dispersal and isolation 

Th eir present world distribution of littoral “terrestrial” arthropods such as geophilomorphs, 

with often widespread genera and species, their occurrence on isolated islands 

and their presence on both sides of oceans could be accounted for by a number of possible 

explanations including continental drift, climate change, transport by birds or 

other animals aerial dispersal, human activity and passive transport by water. 

With Neotropical geophilomorphs there are suggestions that a few taxa have the 

traits of an old Gondwanan faunal element but the bulk of the group belong to wideranging 

groups, possibly recent immigrants to South America (Pereira et al., 1997 ). 

Verhoeff ( 1935 ) when fi rst describing Strigamia japonica , suggested that there might 

have once been a continuous occurrence of Strigamia along the Siberian coast during 

the last warm period, later broken up by climate change leading to the separation of 

the European and Asian maritime forms. 

Transport by birds has been suggested for certain freshwater millipedes (Golovatch 

and Kime, 2009 ) but there is no defi nite evidence of this being a common mechanism 

for dispersing littoral arthropods. Also small organisms are always at risk of aerial transport 

as dust, etc. is picked up and carried in the atmosphere, possibly for long distances 

but, again, this is probably unlikely for littoral centipedes. Dispersal by human activity 

(anthropochory) is always seen as a way by which animals and plants are spread around 

the world and many animals seem to have been dispersed in this way, as for instance, 

North American invertebrates (Lindroth, 1957 ), Newfoundland centipedes (Palmén,


1954 ) and the myriapods of St.Helena (Matic and Darabantu, 1977 ) all of which 

include European species. Crabill ( 1964 ) thought that Maoriella edentatus (Attems, 

1947) described from Tahiti (no locality details) might possibly be an aberrant 

Maoriella macrostigma Attems 1903 transported from New Zealand by early Polynesian 

voyagers; Maoriella is a genus known otherwise only from New Zealand with one species 

from Australia. “Terrestrial” (especially soil) animals, especially those associated 

with agriculture or similar practices, would seem to be likely to be spread this way 

rather than sea shore organisms although the possibility of the latter being carried in 

ship’s ballast should not be forgotten. 

Accidental dispersal by rafting of animals e.g. on plant debris (hydrochory) is certainly 

seen as a likely dispersal mechanism, for animals and littoral species are in an 

optimum situation for this, both in terms of being accidentally carried away and of 

establishing themselves when they arrive at a suitable destination. Suomalainen ( 1939 ) 

had reported on Pachymerium ferrugineum fl oating on sea water for as long as 31 days 

before sinking and long survival when submerged in such water. Anecdotally, Nesrine 

Akkari ( pers. comm .) described how a live Pachymerium ferrugineum was taken directly 

in a landing net from the water in a shallow lagoon in Tunisia by a colleague who was 

collecting mosquito larvae. Interestingly, this species is probably tone of the most widespread 

centipedes in the world. Fossil evidence suggests that rafting occurred in palaeooceans 

and during recent centuries the composition and abundance of fl oating items 

has been strongly aff ected by human activities; it is suggested that rafting continues to 

be an important dispersal in present-day oceans (Th iel and Gutow, 2004 ). 

Th ere is, in fact, very limited quantitative data on the extent to which even insects 

actually survive as drifters and most studies on animals spreading by rafting concentrate 

on marine species (e.g. Th iel and Gutow, 2005 ). Drifting vegetation has been 

found up to 16km off shore which supported many insects although only 25% of it had 

living terrestrial animals and no living insects were found in vegetation collected 160km 

off shore (although colonies of ants have been reported in drifting wood) (Bowden and 

Johnson, 1976 ). Even if the chances of survival are small, the possibility remains for 

transport in this way. Th e pseudoscorpion Apocheiridium pelagicum Redikorzev, 1938 

was fi rst collected 200 miles (320km) at sea in plankton nets; its habitat is in reefs 

constantly submerged by the sea (Roth and Brown, 1976 ). Th e fact that heteropterous 

bugs ( Halobates spp.) can live on the surface of the open ocean suggests also that there 

is no fundamental reason why arthropods of terrestrial origin should not be able to 

survive in this environment for a period of time and be dispersed over wide distances. 

Crabill ( 1960 ) discussed the issue of rafting as a means of distribution of centipedes 

referring to Caritohallex minirrhopus , Balophilus riveroi and Schendylus virgingordae collected 

in West Indian beach drift and to specimens of Pectiniunguis from seaweed on a 

beach in Florida Key. He thought that animals might not be “discomfi ted much or at 

all perhaps” in the wood or under the bark of fl oating trees or bushes and suggesting 

that, although many, perhaps the majority, would perish, over the immense length of 

time and the millions of such voyages that were begun, many must have reached land 

and survived. He went on to comment that “during the immense stretches of time of 

the past some centipedes – they may well have been Schendylidae – made the journey


successfully and this explanation reasonably accounts for the presence in Africa and 

South America of many, but of course not all, congeneric or conspecifi c centipedes”. 

Th is hypothesis was supported by Pereira and Minelli ( 1993 ) and Pereira et al. ( 1997 ), 

the latter writing “Th ere is a single specimen of Schendylops in the Natural History 

Museum in London, too damaged for allow for confi dent specifi c identifi cation, but 

good enough to allow a confi dent identifi cation as a member of a genus occurring, 

with many species, on both sides of the Atlantic. Th is specimen was collected long ago 

from Ascension Island, midway between Africa and South America, along the route 

of the westbound South Equatorial Current”. 

On both a local and a global scale, the varied nature of coasts will be likely to break 

up species into a series of isolated populations which would favour genetic divergence. 

Dispersal across oceans and to isolated islands would accentuate this eff ect. Th is may 

be refl ected in variations in characters between populations at diff erent sites. 

Geophilomorphs are convenient animals in which to study such diff erences by looking 

at variation of numbers of leg-bearing segments at particular locations as described by 

Lewis ( 1962 ) for Strigamia maritima and Shinohara ( 1961 ) for Tuoba littoralis . Arthur 

and Kettle ( 2000 ) demonstrated a latitudinal cline in segment number in Strigamia 

maritima in Britain, suggesting that climatic selection and local adaptation could be 

responsible; Vedel et al. ( 2008 ) showed that, in laboratory experiments, temperature 

regime infl uences the number of leg-bearing segments that develop in this species and 

which could provide an explanation for this phenomenon. 

Conclusions 

It can be seen that there is a wide diversity of species of littoral centipedes around the 

world but with obvious and sometimes surprising gaps in our knowledge including the 

shortage of records from the coasts of eastern North America, most of Africa including 

South Africa and large parts of Asia. Th ere are also littoral centipedes, presumably 

geophilomorphs, reported in the literature from a variety of locations when, apparently, 

no identifi cation was made or specimens kept e.g. Cape Verde Islands, Galapagos 

(Crossland, 1929 ), Colombia, Panama, Costa Rica (Polhemus and Evans, 1969 ), 

North America (Hoff man and Carlton, 2007 ). Th ere may also be described species 

that have been missed in this survey. 

On northwestern European coasts alone there are at least eight or so genuini or indifferenti 

species described (depending on taxonomic issues, etc.) and the number from 

Japanese shores is comparable. It would be surprising if such relative species richness 

was not replicated in at least some other regions and that areas from which only one or 

two or no species have been described are probably in many cases much under recorded. 

It seems highly likely, therefore, that there are further, as yet undescribed, littoral 

geophilomorphs and/or a wider distribution of known species than has already been 

recorded. 

What we have is a somewhat incomplete picture but one that suggests that, wherever 

conditions are favourable (outside the polar regions), seashore geophilomorphs are


likely to be found on coastlines around the world. Furthermore, their apparent tolerance 

of seawater would facilitate dispersal e.g. by rafting across oceans and a widespread 

distribution may be the eff ect of this rather than by accidental human transmission. 

What is very noticeable is that littoral species are not confi ned to one genus or family; 

some are in monotypic genera, others in genera with many species. Table 3 lists 

genera with numbers of valid species listed in Chilobase ( http://chilobase.bio.unipd.it ) 

and numbers of halophilic species. For all the reasons already mentioned, numbers are 

only approximate and, in addition, defi nitions/descriptions of genera and species are 

open to revision but this gives us a crude idea of the relative numbers of species in 

diff erent genera. 

Divergence between the centipede orders seems to date from the Silurian and early 

Devonian with familial divergences also almost wholly Palaeozoic (Murienne et al., 

2010). As modern families, genera and species have evolved certain forms in some of 

these have come to inhabit halophilic habitats either in addition to their more terrestrial 

ones or on an apparently obligate basis. As well as the monotypic genera (which 

would be derived from terrestrial ones) and doubtful halophiles, a dozen genera include 

one or more halophilic species suggesting that, within each of these genera, certain 

species have become ecologically specialised as inhabitants of the littoral zone. If this is 

the case, and assuming that this has happened independently in each genus (possibly 

more than once), then the adoption of the halophilic habit by some geophilomorphs 

Table 3. Genera with known halophilic species, and their relative number. 

Family Genus Number of species 

in genus 

Halophile 

species (s. l.) 

Percentage 

Mecistocephalidae Mecistocephalus 131 2 1.5 

Schendylidae Hydroschendyla 1 1 monotypic 

Nyctunguis 17 1 5.9 

Pectiniunguis 23 6 26.1 

Schendyla 27 2 7.4 

Schendylops 58 1 1.7 

Th indyla 1 1 monotypic 

Ballophilidae Caritohallex 1 1 monotypic 

Ballophilus 36 1 2.8 

Lintotaeniidae Strigamia 39 2 5.1 

Dignathodontidae Henia 18 1 5.6 

Geophilidae Erithophilus 1 1 monotypic 

Geophilus 136 7 5.1 

Lionyx 1 1 monotypic 

Maoriella 6 1 16.7 

Mixophilus 1 1 monotypic 

Pachymerium 22 1 4.5 

Polycricus 23 1 4.3 

Tuoba 17 11 64.7 

Tretecthus 1 1 monotypic 

Aphilodontidae Aphilodon 14 ?1 ?7.1


must have happened on a number of occasions since the order fi rst appeared. 

Th is certainly suggests features of the group that, in some way, “pre-adapts” them 

for this habitat without, apparently, a need for a great deal of morphological (and presumably 

physiological) specialisation. In some genera, especially large ones such as in 

the genus Geophilus this colonisation of the seashore could have happened several 

times. In the case of miriopodi indiff erenti there seems to be no real obstacles to them 

occupying niches in both truly terrestrial as well as upper shore habitats. For the genuin 

i, presumably greater specialisation or perhaps competition confi nes them to the 

littoral habitat. Maybe for those forms apparently halofi li genuini in some areas but 

terrestrial in others perhaps we should look for either specifi c local factors, possible 

population diff erences/isolation, taxonomic issues or simply lack of adequate distribution 

and habitat knowledge. 

Two genera, Pectiniunguis and Tuoba appear to have a relatively high proportion of 

halophilic species and, interestingly, these are the ones identifi ed as having the distinctive 

pre-tarsal parunguis. Tuoba has often been considered as largely littoral genus 

(Lewis, 1996 ) and of the 17 species described, nearly two-thirds are littoral. Th e 

Australian Tuoba sydneyensis seems to have reached Hawai’i (possibly on drifting material) 

and maybe has been able to colonise otherwise presumably vacant niches and 

reversed the trend, going from essentially littoral to terrestrial. It could well be that 

other species have done the same. 

Acknowledgements 

Th e pioneering work of John Lewis both on littoral geophilomorphs and in his 1981 

book set the scene from which this account could be built up using a diversity of 

sources and from many individuals, most of whom are acknowledged in the earlier 

paper (Barber, 2009a ). To these I would like to add Matty Berg, Martin Cawley, Andrey 

Przhiboro, Randy Mercurio and Irina Zenkova. Also the several unknown reviewers 

who read the fi rst draft of this paper and most helpfully pointed out various errors of 

emphasis, interpretation and typography. 

References 

Andersson , G . 1983 . Th e Chilopod fauna in the vicinity of Göteborg – a comparison between collecting 

results obtained in the 1920s and the 1970s . Acta Entomologica Fennica 42 : 9 - 14 

Andersson , G. , B. A. Meidell , U. Scheller , J.-Å. Winquist , M. Osterkamp Madsen , P. Djursvol , G. Budd , 

and U. Gärdenfors . 2005 . Nationalnyckeln till Sveriges fl ora och fauna. Mångfotingar. Myriapoda. 

ArtDatabanken, SLU . Uppsala , Sweden . 351 pp. 

Armitage , P. 1982 . Strigamia maritima (Leach) (Chilopoda, Geophilomorpha), fi rst record in fresh water. 

Entomologists’ Monthly Magagazine 118 : 43 - 44 . 

Arthur , W. and C. Kettle . 2000 . Latitudinal cline in segment number in an arthropod species, Strigamia 

maritima . Proceedings of the Royal Society of London B 267 : 1393 - 1397 . 

Barber , A. D . 2009a . Littoral myriapods; a review in Myriapoda and Onychophora of the World – 

Diversity, biology and importance . Xylander, W and K. Voigtländer (Editors). Soil Organisms 81 (3) : 

735 - 760 .


Barber , A. D . 2009b . Centipedes. Linnean Society Synopses of the British Fauna (NS) 58. Field Studies 

Council. Shrewsbury , England , UK . 228 pp. 

Berg , M. P. , M. Soesbergen , D. Templeman , and H. Wijnhoven . 2008 . Verspreidingsatlas Neder landes 

landpissebedden, duizendpoten en miljoenpoten (Isopoda, Chilopoda, Diplopoda). Vrije Universiteit, 

Afdeling Dierecologie. Leiden , EIS-Nederland and Amsterdam , Th e Netherlands . 192 pp. 

Bergersen , R. , K. M. Olsen , P. Djursvol , and A. Nilssen . 2006 . Centipedes (Chilopoda) and millipedes 

(Diplopoda) in North Norway . Norwegian Journal of Entomology 53 : 23 - 38 . 

Binyon , J. and J. G. E. Lewis . 1963 . Physiological adaptations of two species of centipede (Chilopoda: 

Geophilomorpha) to life on the shore . Journal of the Marine Biological Association of the United 

Kingdom 43 : 49 - 55 . 

Bonnel , B . 1929 . Geophilid centipedes from the bed of the Coum River (Madras) . Journal of the Asiatic 

Society of Bengal (New Series) 25 : 181 - 4 . 

Bowden , J. and C. G. Johnson . 1976 . Other intertidal air-breathing arthropods. Migrating and other terrestrial 

insects at sea . Chapter 6, pp. 97-118 . In: Marine Insects. Cheng , L. (ed.). North Holland 

Publishing , Amsterdam. Th e Netherlands . 581 pp. 

Brölemann , H. W. 1930 . Éleménts d’une faune de France Chilopodes . Lechavalier , Paris, France . 

405 pp. Also available at : http://www.faunedefrance.org/bibliotheque/docs/H.W.BROLEMANN 

(FdeFr25)Myriapodes-Chilopodes.pdf . 

Catenazzi , A. and M. A. Donnelly . 2007 . Th e Ulva connection: marine algae subsidize terrestrial predators 

in coastal Peru . Oikos 116 : 75 - 86 . 

Chamberlin , R. V . 1909 . Some records of North American Geophilidae and Lithobiidae. With description 

of new species . Annals of the Entomological Society of America 11 (3) : 175 - 192 . 

Chamberlin , R. V . 1960 . Five new western geophilid chilopods . Proceedings of the Biological Society of 

Washington 73 : 239 - 244 . 

Cheng , L . 1976 . Marine Insects . North Holland Publishing , Amsterdam. Th e Netherlands . 581 pp. 

ChiloBase. (No date) A world catalogue of centipedes on line (A. Minelli, Editor) http://chilobase 

.bio.unipd.it/ Version 1.01 was dated 2006 but this database is being continuously updated. Note 

for the reader: if opening this link directly does not seem to work, simply google the word 

“Chilobase”. 

Crabill, R. E. Jr . 1960 . Centipedes of the Smithsonian-Bredin expeditions to the West Indies . Proceedings 

of the United States National Museum 111 : 167 - 195 . 

Crabill, R. E. Jr . 1964 . A preliminary review of Maoriella , with description of a new species from the 

Chatham Islands (Chilopoda: Geophilomorpha: Chilenophilidae) . Entomological News 75 (4) : 

85 - 97 . 

Crabill, R. E. Jr . 1968 . On the true identities of Tuoba and Nesogeophilus (Chilopoda: Geophilomorpha: 

Geophilidae) . Proceedings of the Entomological Society of Washington 70 : 345 . 

Crossland , C . 1929 . Amphibious Centipedes . Nature 124 : 794 . 

Due , A. D. and G. A. Polis . 1985 . Th e biology of Vaejovis littoralis Williams, an intertidal scorpion from 

Baja California, Mexico . Journal of Zoology London (A) 207 : 563 - 580 . 

Evans , W. G. 1980 . Insects, Chilopoda and Arachnida: Insects and allies . pp. 641-658. In , Morris, R. H. , 

D. P. Abbott , and E. C. Haderlie . Th e Intertidal Animals of California . Stanford University Press . 

Stanford, California, USA . 690 pp. + xi . 

Geoff roy , J-J. and E. Iori . 2009 . Th e French centipede fauna (Chilopoda): updated checklist and distribution 

in mainland France, Corsica and Monaco . In: Myriapoda and Onychophora of the World – 

Diversity, biology and importance ( Xylander, W and K. Voigtländer , eds). Soil Organisms 

81 (3) : 671 - 694 . 

Golovatch , S. I. and R. D. Kime . 2009 . Millipede (Diplopoda) distributions: A review . In: Myriapoda and 

Onychophora of the World – Diversity, biology and importance ( Xylander, W and K. Voigtländer , 

eds). Soil Organisms 81 (3) : 565 - 597 . 

Haberman , K. L . 1982 . Vertical distribution and habitat of the centipede Geophilus becki in the high 

intertidal and splash zone of Monterey Bay , California . Hopkins Marine Station student paper : Final 

papers biology 175H (unpublished).


Harger , O . 1872 . Description of New North American Myriapods . American Journal of Science and Arts 

(Ser. 3) 4 : 116 - 121 . 

Hennings , C . 1903 . Zur Biologie der Myriopoden I Marine Myriopoden . Biologisches Zentralblatt 23 : 

720 - 725 . 

Hoff man , R. and J. T. Carlton . 2007 . Chilopoda pp. 692-693 . In: Light, S.F. and J. T. Carlton. Light & 

Smith Manual: Intertidal invertebrates from Central California to Oregon . Fourth edition . University 

of California Press. Berkley, California , USA . 1001 pp. 

Hoback , W. W. and D. W. Stanley . 2001 . Insects in hypoxia . Journal of Insect Physiology 47 (6) : 

533 - 542 . 

Horneland , E. O. and B. A. Meidell . 1986 . Th e epimorphosis of Strigamia maritima (Leach, 1817) 

(Chilopoda, Geophilidae) . Entomologica Scandinavica 17 : 127 - 129 . 

Iorio , E . 2005 . Contribution à la connaissance des chilopodes de Bretagne (Myriapoda, Chilopoda) . 

Bulletin du Société Linnéanne du Bordeaux 140 (33) : 149 - 156 . 

Ivinskis , P. and J. Rimšaitė . 2005 . Baltic seashore as a unique habitat for insects . Acta Zoologica Lituanica 

15 (2) : 115 - 118 . 

Laloy , L . 1904 . Insectes arachnides et myriapods marines . La Nature (Paris) 132 : 154 - 155 . 

Leach , W.E . ( 1817 ) Th e Characters of the Genera of the Class Myriapoda, with Descriptions of some 

Species . Zoological Miscellany 3 (12) : 31 - 45 . 

Lewis , J. G. E. 1960. Th e life history and ecology of the littoral centipede Strigamia maritima (Leach). 

PhD Th esis. University of London (unpublished) . Cited in Lewis , J.G.E . ( 1981 ). 

Lewis , J. G. E . 1961 . Th e life history and ecology of the littoral centipede Strigamia (= Scolioplanes ) maritima 

(Leach) . Proceedings of the Zoological Society of London 117 (2) : 221 - 248. 

Lewis , J. G. E . 1962 . Th e ecology, distribution and taxonomy of the centipedes found on the shore in the 

Plymouth area . Journal of the Marine Biological Association of the United Kingdom 42 : 655 - 664 . 

Lewis , J. G. E . 1981 . Th e biology of centipedes . Cambridge University Press . Cambridge, England, UK . 

476 pp. 

Lewis , J. G. E . 1996 . On a new species of Tuoba from Ascension Island . Fragmenta Entomologica 28 

(1) : 15 - 20 . 

Lindroth , C. H . 1957 . Th e Faunal connections between Europe and North America. Wiley / Almqqvist 

& Wiskell . New York, NY, USA / Stockholm , Sweden . 326 pp. 

Little , C . 1983 . Th e colonisation of land: origins and adaptations of terrestrial animals . Cambridge 

University Press . Cambridge, England, UK . 299 pp. 

Little , C . 1990 . Th e terrestrial invasion, An ecophysiological approach to the origins of land animals . 

Cambridge University Press. Cambridge, England, UK . 304 pp. 

Lock , K . 2009 . Updated checklist of the Belgian centipedes (Chilopoda) . Entomologie faunistique – 

Faunistic Entomology 62 : (1) : 35 - 39 . 

McQueen , D. J. and C. L. McLay . 1983 . How does the intertidal spider Desis marina (Hector) remain 

under water for such a long time? New Zealand Journal of Zoology 10 (4) : 383 - 392 . 

Manton , S. M . 1965 . Th e evolution of arthropodan locomotory mechanisms 8. Functional requirements 

and body design in Chilopoda, together with a comparative account of their skeleton-muscular systems 

and an Appendix on A comparison between burrowing forces of annelids and chilopods and its 

bearing on the evolution of the arthropodan haemocoel . Journal of the Linnean of London (Zoology ) 

46 (306-7) : 251 - 483 . 

Matic , Z. and C. Darabantu . 1977 . La faune terrestre de l’Îsle de Sainte-Hélène, Quatrième Partie. Musee 

Royal de l’Afrique Centrale,Tervuren, Belgique. Annales-Serie IN-8 o , Sciences Zoologiques 220 : 

45 - 359 . 

Meidell , B. A . 1979 . Norwegian Myriapoda: Some Zoogeographical Remarks . pp 195-201 . In: 

Camatini, M. ( ed.). Myriapod Biology. London, England , UK. Academic Press . 456 pp. 

Morse , D. H . 1997 . Distribution, movement, and activity patterns of an intertidal wolf spider Pardosa 

lapidicina population (Araneae, Lycosidae) . Journal of Arachnolog y 25 : 1 - 10 . 

Mundel , P. , 1979 . Th e centipedes (Chilopoda) of the Mazon Creek . pp. 361–378 . In: Nitecki, M. H. 

(ed.). Mazon Creek Fossils . Academic Press . New York, NY, USA . 581 pp.


Murienne , J. , G. D. Edgecombe , and G. Giribet . 2010 . Including secondary structure, fossils and 

molecular dating in the centipede tree of life . Molecular Phylogenetics and Evolution 57 : 301 - 313 . 

Palmén , E . 1949 . Th e Chilopoda of Eastern Fennoscandia. Annales Zoologicae Societatis . Zoologicae 

Botanicae Fennicae ‘Vanamo’ 13 (4) : 1 - 45 . 

Palmén , E . 1954 . Survey of the Chilopoda of Newfoundland. Archivum Societatis . Zoologicae Botanicae 

Fennicae ‘Vanamo’ 8 (2) : 131 - 149 . 

Palmén , E. and M. Rantala . 1954 . On the life history and ecology of Pachymerium ferrugineum (C.L.Koch) 

(Chilopoda, Geophilidae). Annales Zoologice Societatis . Zoologicae Botanicae Fennicae ‘Vanamo’ 

16 (3) : 1 - 44 . 

Pearse , V. J. , M. Pearse , M. Buchsbaum , and R. Buchsbaum , R. 1987 . Living Invertebrates . 

Blackwell Scientifi c Press . ( Palo Alto, California, USA ) & Boxwood Press (Pacifi c Grove, California) . 

862 pp. 

Pereira , L. A. , D. Foddai, and A. Minelli . 1997 . Zoogeographical aspects of Neotropical Geophilomorpha 

(Chilopoda) . Entomologica Scandinavica Supplement 51 : 77 - 86 . 

Pereira , L. A. , D. Foddai and A. Minelli . 1999 . Pectiniunguis bollmani n.sp. from Corraline Island Cayo 

Sombrero (Venezuela) with notes on P.hallirhytus Crabill, 1959 (Chilopoda: Geophilomorpha: 

Schendylidae) . Studies in Neotropical Fauna & Environment 34 ( 3 ): 176 - 185 . 

Pereira , L. A. and A. Minelli , A. 1993 . On two new species of Schendylurus Silvestri 1907 from Venezuela, 

with redescription of S. colombianus Chamberlin 1921 and S . virgingordae Crabill 1960 (Chilopoda 

Geophilomorpha Schendylidae). Tropical Zoology , Special Issue 1 : 105 - 123 

Pisani , D. , L. L. Poling , M. Lyons-Weiler , S. B. Hedges . 2004 . Th e colonisation of land by animals: 

molecular phylogeny and divergence times among arthropods . BMC Evolutionary Biology 2 : 1 - 10 . 

Plateau , F . 1890 . Les myriopodes marins et la résistance des arthropods a respiration aérienne a la submersion 

. Journal de l’anatomie et physiologie normales et pathologiques de l’hommes et des animaux 

26 : 236 - 269 . 

Polhemus , J. T. and W. G. Evans . 1969 . A new genus of intertidal Saldidae from the Eastern tropical 

Pacifi c with notes on its biology (Hemiptera) . Pacifi c Insects 11 (3-4) : 571 - 578 . 

Polis , G. A. and S. D. Hurd . 1995 . Extraordinarily high spider densities on islands: Flow of energy from 

the marine to terrestrial food webs and the absence of predation . Proceedings of the National 

Academy of Sciences of the United States 92 (10) : 4382 - 4386. 

Polis , G. A. , M. E. Power , and G. R. Huxel . (Editors) 2004 . Food webs at the landscape level . University 

of Chicago Press . Chicago, Illinois, USA . 548 + ix pp. 

Przhiboro , A. 1994 . Fauna of terrestrial arthropods of the intertidal zone of Kandalaksha Bay, the White 

Sea . Unpublished M. Sc . Th esis. St. Petersburg State University . St. Petersburg, Russia . 128 pp. + 128 

pp. of supplementary material . 

Rajalu , G. S . 1970 . Tracheal pulsation in a marine centipede Mixophilus indicus . Current Science (Indian 

Academy of Sciences) 17 : 397 - 398 . 

Rajalu , G. S . 1972 . On the mode of respiration of an estuarine centipede Mixophilus indicus . Journal of 

Animal Morphology and Physiology 9 : 181 - 90 . 

Ricketts , E. F. , J. Calvin , and J. W. Hedgepeth . 1985 . Between Pacifi c Tides . 5 th Edition . Revised by 

D. W. Phillips . Stanford University Press, Stanford , California, USA . pp. 652 . 

Roth , V. D. and W. L. Brown . 1975 . A new genus of Mexican intertidal zone spider (Desidae) with biological 

and behavioural notes . American Museum Novitates 2568 : 1 - 7 . 

Roth , V. D. and W. L. Brown . 1976 . Other Intertidal air-breathing arthropods . Chapter 6 , pp. 119-150 

In: Cheng , L. (ed.). North Holland Publishing , Amsterdam. Th e Netherlands . 581 pp. 

Rosenberg , J . 2009 . Der Hundertfüßer . Die Neue Brehm-Bücherei 285 . Westarp Wissensschaften, 

Hohenwarsleben , Germany . 524 pp. 

Rovner , J. S . 1987 . Nests of terrestrial spiders maintain a physical gill: fl ooding and the evolution of silk 

constructions . Journal of Arachnology 14 : 327 - 337 . 

Schubart , O . 1929 . Th alassobionte und thalassophile Myriapoda . In: Grimpe, G. and E. Wagler (eds). 

Tierwelt der Nord- und Ostsee 11 : 1 - 20 .


Schmitz , A. and J. F. Harrison . 2004 . Hypoxic tolerance in air-breathing invertebrates . Respiratory 

Physiology & Neurobiology 141 (3) : 229 - 242 . 

Selden , P. A. and D. Edwards . 1989 . Colonisation of the land . Chapter 6. pp. 122-152. In: Allen, K. C. 

and D. E. G. Briggs (eds). Evolution and the fossil record . Bellhaven Press , London, England, UK & 

Smithsonian Institution Press, Washington, District of Columbia, USA. xiii + 265 pp. 

Shear , W. A. and G. D. Edgecombe . 2010 . Th e geological record and phylogeny of the Myriapoda . 

Arthropod Structure & Development 39 : 174 - 190 . 

Shear , W. A. and P. A. Selden . 2001 . Rustling in the undergrowth; animals in early terrestrial ecosystems 

pp. 29-51. In: Gensel, P.A. and D. Edwards. Plants invade the land . Columbia University Press . New 

York, NY, USA . 304 pp. 

Shinohara , K . ( 1961 ) Survey of the Chilopoda and Diplopoda of Manazuru Seashore, Kanagawa 

Prefecture, Japan . Science Reports Yokosuka City Museum 6 : 75 - 82 . 

Silvestri , F . 1903 . Miriopodi viventi sulla spiaggia del mare presso portici (Napoli) . Annuari del Museo 

Zoologico della Universita di Napoli (Nuova Serie) 1 (12) : 1 - 5 . 

Silvestri , F . 1929 . Description of a new genus and species of Geophilidae (Myriapoda, Chilopoda) from 

Madras (India) . Records of the Indian Museum (Calcutta ) 31 : 263 - 267 . 

Simiakis , S. , A. Minelli , and M. Mylonas . 2004 . Th e centipede fauna (Chilopoda) of Crete and its 

satellite islands (Greece, Eastern Mediterranean) . Israel Journal of Zoology 50 : 367 - 418 . 

Suomalainen , P . 1939 . Zur Verbreitungsökologie von Pachymerium ferrugineum C.Koch (Myriopoda) in 

fi nnischen Schärenhof . Annales Zoologici Societatis Zoologicae Botanicae Fennicae ‘Vanamo’ 7 : 

10 - 14 . 

Th iel , M. and L. Gutow , L. 2004 . Th e ecology of rafting in the marine environment I: Th e fl oating substrata 

. Oceanography & Marine Biology 42 : 181 - 264 . 

Th iel , M. and L. Gutow . 2005 . Th e ecology of rafting in the marine environment II: Th e rafting organisms 

and community . Oceanography & Marine Biology 43 : 279 - 418 . 

Uliana , M. , L. Bonato , and A. Minelli . 2007 . Th e Mecistocephalidae of the Japanese and Taiwanese 

Islands ( Chilopoda : Geophilomorpha ). Zootaxa 1396 : 1 - 84. 

Vedel , V. , A. D. Chipman , M. Akam , and W. Arthur . 2008 . Temperature-dependent plasticity of segment 

number in an arthropod species: the centipede Strigamia maritima . Evolution & Development 

10 (4) : 487 - 492. 

Verhoeff , K . 1935 . Über Scolipolanes (Chilopoda) . Zoologischer Anzeiger 111 : 10 - 23 . 

Wilson , H. M. and L. I. Anderson . 2004 . Morphology and taxonomy of Paleozoic millipedes (Diplopoda: 

Chilognatha: Archipolypoda) from Scotland . Journal of Paleontology 78 : 169 - 184 .

Geophilomorph centipedes and the littoral habitat - Books and ...

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?