Gonggok atau juga dikenali sebagai ulat geluntur, sepah bulan atau mentibangmerupakan sejenis artropod dengan badan

berbentuk silinder yang panjang. Badan gonggok mempunyai rangka luar (exoskeleton) dan terbahagi kepada antara 20 hingga 100 segmen yang bersambung sambung. !etiap segmen mempunyai dua pasang kaki(ke"uali segmen pertama selepas kepala yang tiada kaki langsung, dan beberapa segmen selepasnya yang mempunyai hanya sepasang kaki). #erdapat kira kira 12,000 jenis spesies gonggok dalam kira kira 1$0 %amili.&1' Gonggok berjalan. ( Gonggok menggulungkan diri apabila diganggu

#emporal range) $2*+0,a -re. . / ! 0 1 - # 2 3 -g 4 5ate !ilurian to 6e"ent 6usty millipede (#rigoniulus "orallinus) !"ienti%i" "lassi%i"ation 3ingdom 7nimalia ) -hylum) 7rthropoda

!ubphylu m) 1lass)

,yriapoda

0iplopoda 0e Blain8ille in Ger8ais, 1*$$ !ub"lasses ( -eni"illata ( 1hilognatha ( 7rthropleuridea (extin"t) 0i8ersity 19 orders, ". 12,000 spe"ies

,illipedes ("lass 0iplopoda) are myriapodous arthropods that ha8e t:o pairs o% legson most body segments. ;a"h double legged segment is a result o% t:o single segments %used together as one (the name <0iplopoda< "omes %rom the Greek :ords=>?@AB (diplous), <double< and ?A=CB (podos), <%oot<). ,ost millipedes ha8e 8ery elongated "ylindri"al or %lattened bodies :ith more than 20 segments, :hile pill millipedes are shorter and "an roll into a ball, like a pillbug. #he name <millipede< is a "ompound :ord %ormed %rom the 5atin roots mille(<thousand<) and pes (<%oot<). 0espite their name, no kno:n millipede has 1,000 legs, although the rare spe"ies Dlla"me plenipes has up to EF0.&1' 1ommon spe"ies ha8e bet:een G9 and $00 legs. #here are approximately 12,000 named spe"ies in "a. 1$0 %amilies.&2' #he longest spe"ies is the giant 7%ri"an millipede (7r"hispirostreptus gigas). ,ost millipedes are slo: mo8ing detriti8ores, eating de"aying lea8es and other deadplant matter. Ho:e8er, they "an also be minor garden pests, espe"ially in greenhouses:here they "an "ause se8ere damage to emergent seedlings. ,illipedes "an be easily distinguished %rom the some:hat similar and related"entipedes (1lass 1hilopoda) :hi"h mo8e rapidly, are "arni8orous, and ha8e a single pair o% legs %or ea"h body segment. #he s"ienti%i" study o% millipedes is kno:n asdiplopodology, and a s"ientist :ho studies them is "alled a diplopodologist. 1ontents &hide' ( 1 ;8olution 2 1hara"teristi"s ( 2.1 Head ( 2.2 Body ( 2.G Dnternal organs ( G 6eprodu"tion and gro:th $ ;"ology

( $.1 Habitat ( $.2 0iet ( $.G -redators and parasites ( $.$ 0e%en"e me"hanisms ( F ,illipedes in relation to people ( 9 1lassi%i"ation ( E !ee also ( * 6e%eren"es ( I ;xternal links ;8olution&edit' #his "lass o% arthropod is thought to be among the %irst animals to ha8e "olonised land during the !ilurian geologi" period. #hese early %orms probably ate mosses and primiti8e 8as"ular plants. #he oldest kno:n land "reature, -neumodesmus ne:mani, :as a 1 "m (0.GI in) long millipede, and li8ed $2* million years ago.&G' Dn the Jpper 1arboni%erous (G$0 to 2*0 million years ago),7rthropleura be"ame the largest kno:n land in8ertebrate o% all time, rea"hing lengths o% up to 2.9 m (* %t 9 in). ,illipedes, "entipedes, and other terrestrial arthropods attained 8ery large siKes in "omparison to modern spe"ies in the oxygen ri"h en8ironments o% the 0e8onian and 1arboni%erous periods, and some "ould gro: larger than one metre. 7s oxygen le8els lo:ered through time, arthropods be"ame smaller in siKe.&$' 1hara"teristi"s&edit' #he 4orth 7meri"an millipede 4ar"eus ameri"anus + head :ith eyes, antennae, and enlarged "ollum ,illipede anterior anatomy ,illipedes range %rom 2 mm (0.0EI in) to around GF "m (1$ in) in length,&F' and "an ha8e as %e: as ele8en to o8er one hundred segments. #hey are generally bla"k or bro:n in "olour, although there are a %e: brightly "oloured spe"ies. #he millipedeLs most ob8ious %eature is its large number o% legs. Ha8ing many short legs makes millipedes rather slo:, but the many legs pushing in unison pro8ides po:er%ul strength %or burro:ing. Head&edit' #he head o% a millipede is typi"ally rounded abo8e and %lattened belo: and bears a pair

o% large mandibles in %ront o% a plate like stru"ture "alled a gnatho"hilarium (<ja: lip<).&9' #he head "ontains a single pair o% antennae :ith se8en or eight segments and a group o% sensory "ones at the tip.&9' ,any orders also possess a pair o% sensory organs kno:n as the #MmMs8Nry organs, shaped as small o8al rings posterior and lateral to the base o% the antennae. #heir true %un"tion is unkno:n,&9' but they also o""ur in some "entipedes, and are possibly used to measure humidity or light le8els in the surrounding en8ironment. &E' ,illipede eyes "onsist o% a number o% simple %lat lensed o"elliarranged in a group on the side o% the head. ,any spe"ies o% millipedes, in"luding the entire order -olydesmida and "a8e d:elling millipedes su"h as 1auseyella and #ri"hopetalum, ha8e se"ondarily lost their eyes and are "ompletely blind.&F' Body&edit' -aranota o% a -olydesmidan (le%t) and-latydesmidan millipede #he body is %lattened or "ylindri"al, "omposed o% numerousmetemeri" segments, ea"h :ith an exoskeleton "onsisting o% a single "hitinous plate abo8e, one at ea"h side, and t:o or three on the underside. Dn many millipedes, these plates are %used to 8arying degrees, sometimes %orming a single "ylindri"al ring. #he plates are typi"ally hard, being impregnated :ith "al"ium salts.&*' Be"ause they la"k a :axy "uti"le, millipedes are sus"eptible to :ater loss and must spend most o% their time in moist or humid en8ironments.&I' #he %irst segment behind the head is legless and kno:n as a "ollum. #he se"ond to %ourth body rings bear a single pair o% legs ea"h and are kno:n as <haplosegments< (%rom the Greek haplo) single). #he remaining segments, %rom the %i%th to the posterior, are properly kno:n as diplosegments, or double segments. ;a"h diplosegment bears t:o pairs o% legs, rather than just one as in "entipedes. #his is be"ause ea"h diplosgment is %ormed by the %usion o% t:o embryoni" segments. Dn some millipedes the last %e: segments may be legless. #he terms <segment< or <body ring< are o%ten used inter"hangeably to re%er to both haplo and diplosegments. #he %inal segment bears a telson, :hi"h "onsists o% a legless preanal ring, pair o% anal 8al8es ("loseable plates around the anus), and a subanal s"ale.&9'&*' ,illipedes in se8eral orders ha8e keel like extensions o% the body :all kno:n as paranota, :hi"h "an 8ary :idely in shape and siKe. #hey may allo: millipedes to :edge more se"urely into "re8i"es, prote"t the legs, or make the millipede more di%%i"ult %or predators to s:allo:.&10' Dnternal organs&edit'

,illipedes breathe through t:o pairs o% spira"les lo"ated 8entrally on ea"h segment.&11' ;a"h opens into an internal pou"h, and "onne"ts to a system o% tra"heae. #he heart runs the entire length o% the body, :ith an aorta stret"hing into the head. #he ex"retory organs are t:o pairs o% malpighian tubules, lo"ated near the mid part o% the gut. #he digesti8e tra"t is a simple tube :ith t:o pairs o% sali8ary glands to help digest the %ood. &*' 6eprodu"tion and gro:th&edit' ;pibolus pul"hripes mating ,illipedes sho: a di8ersity o% mating styles and stru"tures. Dn the basal order-olyxenida, males deposit spermatophores that are subseOuently pi"ked up by %emales.&11' Dn all other millipede groups, males possess one or t:o pairs o% modi%ied legs "alled gonopods :hi"h are used to trans%er sperm to the %emale during "opulation. #he lo"ation o% the gonopods di%%ers bet:een groups) in males o% the-entaKonia they are lo"ated at the rear o% the body and kno:n as telopods, :hile in Helminthomorpha + the 8ast majority o% spe"ies + they are lo"ated on the Eth body segment.&9' 7 %e: spe"ies are parthenogeneti", ha8ing %e:, i% any, males.&12'&1G' Gonopod morphology o% /xidus gra"ilis(-aradoxosomatidae). Palse "olor !;,image, s"ale bar) 0.2 mm Gro:th stages o% 4emasoma(4emasomatidae), :hi"h rea"hes reprodu"ti8e maturity in stage Q. #he mature gonopods de8elop %rom the legs o% segment E. Gonopod morphology is the predominant means o% determining spe"ies among millipedes) the "omplex stru"tures di%%er mu"h bet:een spe"ies but 8ery little :ithin a spe"ies.&1$' #he genital openings are lo"ated on the third segment, and are a""ompanied in the male by one or t:o penes :hi"h deposit the sperm pa"kets onto the gonopods. Dn the %emale, the genital pores open into paired small sa"s "alled "yphopods or 8ul8ae, :hi"h are "o8ered by a small hood like "o8er, and are used to store the sperm a%ter "opulation.&*' ,illipede sperm is a%lagellate (la"ks a %lagellum), a uniOue trait among myriapods.&9' Pemales lay bet:een ten and three hundred eggs at a time, depending on spe"ies, %ertilising them :ith the stored sperm as they do so. ,any spe"ies simply deposit the eggs on moist soil or organi" detritus, but some "onstru"t nests lined :ith dried%ae"es, and may prote"t the eggs :ithin silk "o"oons.&*' Dn most spe"ies the %emale abandon the eggs a%ter laying but some spe"ies in the orders -latydesmida and!temmiulida pro8ide parental "are %or eggs and young.&11' #he young hat"h a%ter a %e: :eeks, and typi"ally ha8e only three pairs o% legs, %ollo:ed by up to %our legless segments. 7s they gro:, they "ontinually moult, adding %urther

segments and legs as they do so. !ome spe"ies moult :ithin spe"ially prepared "hambers o% soil or silk,&1F' :hi"h they may also use to :ait out dry :eather, and most spe"ies eat the shed exoskeleton a%ter moulting. ,illipedes li8e %rom one to ten years, depending on spe"ies.&*' ;"ology&edit' Habitat&edit' Dn temperate Kones, millipedes are most abundant in moist de"iduous %orests, but they also o""ur in "oni%erous %orests, deserts, "a8es, and alpine e"osystems.&11' !ome spe"ies "an sur8i8e %resh:ater %loods and li8e submerged under:ater %or up to 11 months.&19' &1E' 7 %e: spe"ies o""ur near the seashore and "an sur8i8e in some:hat salty "onditions. &12'&1*' 0iet&edit' ,ost millipedes are detriti8ores and %eed on de"omposing 8egetation, %ae"es, or organi" matter mixed :ith soil. ,illipedes in the order -olyxenida graKe algae %rom bark, and -latydesmida %eed on %ungi.&9' 7 %e: spe"ies are omni8orous or o""asionally "arni8orous, %eeding on inse"ts, "entipedes, earth:orms, or snails&*'&1I' !ome spe"ies ha8e pier"ing mouth parts that allo: them to %eed on plant jui"es.&11' -redators and parasites&edit' ,illipedes are preyed upon by a :ide range o% animals, in"luding 8arious reptiles, amphibians, birds, mammals, and inse"ts.&9',ammalian predators su"h as "oatis and meerkats roll "aptured millipedes on the ground to deplete de%ensi8e se"retions and rub them o%% the body be%ore "onsuming,&20' :hile "ertain poison dart %rogs are belie8ed to in"orporate the toxi" "ompounds o% millipedes into their o:n de%enses.&21' !e8eral in8ertebrates ha8e spe"ialiKed beha8iors or stru"tures to %eed on millipedes, in"luding lar8al glo::orm beetles,&22' -robolomyrmex ants,&2G' "hlamydephorid slugs,&2$' and the preda"eous dung beetle0elto"hilum 8algum.&2F' 4ar"eus ameri"anus "urled in de%ensi8e position -arasites o% millipedes in"lude nematodes, phaeomyiid %lies, and a"antho"ephalans. &9' 0e%en"e me"hanisms&edit' 0ue to their la"k o% speed and their inability to bite or sting, millipedesL primary de%en"e me"hanism is to "url into a tight "oil + prote"ting their deli"ate legs inside an armoured body exterior. ,any spe"ies also emit 8arious poisonous liOuid se"retions through mi"ros"opi" oKopores (also "alled odori%erous or repugantorial glands), along the sides o% their bodies as a se"ondary de%en"e. #hese se"retions may in"ludealkaloids, benKoOuinones, phenols, terpenoids, andRor hydrogen "yanide, among many others. &29'&2E' !ome o% these substan"es are "austi" and "an burn the exoskeleton o%ants and other inse"t predators, and the skin and eyes o% larger predators. -rimates su"h as

"apu"hin monkeys and lemurs ha8e been obser8ed intentionally irritating millipedes in order to rub the "hemi"als on themsel8es to repel mosOuitoes.&2*'&2I'&G0'!ome o% these de%ensi8e "ompounds also sho: anti%ungal a"ti8ity.&G1' #he bristly millipedes (order -olyxenida) la"k both an armoured exoskeleton and odi%erous glands, and instead are "o8ered in numerous bristles that in at least one spe"ies, -olyxenus %as"i"ulatus, deta"h and entangle ants.&G2' ,illipedes in relation to people&edit' ,illipedes do not bite, and their de%ensi8e se"retions are mostly harmless to humans + usually "ausing only minor dis"oloration on the skin + but the se"retions o% some tropi"al spe"ies may "ause pain, it"hing, lo"al erythema, edema, blisters, e"Kema, and o""asionally "ra"ked skin.&GG'&G$'&GF'&G9' ;ye exposures to these se"retions "auses general eye irritation and potentially more se8ere e%%e"ts su"h as "onjun"ti8itis and keratitis.&GE' Pirst aid "onsists o% %lushing the area thoroughly :ith :aterS %urther treatment is aimed at relie8ing the lo"al e%%e"ts. 7 s:arm o% millipedes !ome millipedes are "onsidered household pests, in"luding Tenobolus "arni%ex :hi"h in%ests that"hed roo%s in Dndia.&G*' /ther spe"ies exhibit periodi"al s:arming beha8iour, :hi"h "an result in home in8asions,&GI' "rop damage,&$0' train delays, or e8en train "rashes and derailments.&$1'&$2' !ome o% the larger millipedes in the orders !pirobolida, !pirostreptida, !phaerotheriidaare popular as pets.&$G' !ome spe"ies "ommonly sold or kept in"lude spe"ies o%7r"hispirostreptus, 7phistogoniulus, 4ar"eus, and /rthoporus.&$$' ,illipedes also appear in %olklore and traditional medi"ine around the :orld. ,any "ultures as"ribe millipede a"ti8ity :ith "oming rains.&$F' Dn the Uoruba "ulture o%4igeria, millipedes are used in pregnan"y and business rituals, and "rushed millipedes are used to treat %e8er, :hitlo:, and "on8ulsion in "hildren.&$9' Dn Vambia, smashed millipede pulp is used to treat :ounds, and in the Ba%ia people o% 1ameroon millipede jui"e is used to treat eara"hes.&$F' Dn "ertain Himalayan Bhotiya tribes, dry millipede smoke is used to treat hemorrhoids.&$E' 4ati8e people in ,alyasia use millipede se"retions in poison tipped arro:s.&$F' #he only reported usage o% millipedes as %ood by humans "omes %rom the Bobo people o%Burkina Paso, :ho "onsume boiled, dried millipedes in tomato sau"e.&$*' 1lassi%i"ation&edit' Por more details on this topi", see 5ist o% millipede %amilies. 7pproximate relati8e di8ersity o% extantmillipede orders,

ranging %rom "a. G,F00 spe"ies o% -olydesmida to 2 spe"ies o%!iphoniulida.&2' 7pproximately 12,000 spe"ies o% millipede ha8e been des"ribed, but the true number o% spe"ies on earth has been estimated at up to *0,000.&9' ,illipedes are "onsidered a mega di8erse yet understudied group, :ith in"onsistent taxonomi" e%%ort o8er time.&$I' -olyxenus lagurus, a spe"ies o% the order -olyxenida Glomeris marginata, a pill millipede o% the order Glomerida /"toglena sierra, a spe"ies o% the order-olyKoniida 7nadenobolus monili"ornis, a spe"ies o% the order !pirobolida Harpaphe haydeniana, a spe"ies o% the order -olydesmida #he li8ing members o% the 0iplopoda are di8ided into sixteen orders in t:o sub"lasses.&2' #he basal sub"lass-eni"illata "ontains small spe"ies :hose exoskeleton is not "al"i%ied, and :hi"h are "o8ered in setae or bristles. 7ll other millipedes belong to the sub"lass1hilognatha, "onsisting o% t:o in%ra"lasses) the in%ra"lass -entaKonia"ontaining the short bodied pill millipedes, and the in%ra"lass Helminthomorpha (:orm like millipedes) "ontaining the great majority o% the spe"ies.&F0'&F1' #he higher le8el "lassi%i"ation o% millipedes is presented belo:, based on !hear, 2011,&2' and !hear W ;dge"ombe, 2010&F2' (extin"t groups). 6e"ent "ladisti" and mole"ular studies ha8e "hallenged traditional "lassi%i"ation s"hemes, and in parti"ular the position o% the orders !iphoniulida and -olyKoniida is not yet :ell established.&9'#he pla"ement and positions o% extin"t groups (X) kno:n only %rom %ossils is tentati8e and not %ully

resol8ed.&9'&F2' 1lass 0iplopoda de Blain8ille in Ger8ais, 1*$$ !ub"lass -eni"illata 5atrielle, 1*G1 ( /rder -olyxenida Qerhoe%%, 1IG$ !ub"lass X7rthropleuridea (pla"ed in -eni"illata by some authors)&F2' ( /rder X7rthropleurida Yaterlot, 1IG$ ( /rder X;oarthropleurida !hear W !elden, 1IIF ( /rder X,i"rode"empli"ida !ub"lass 1hilognatha 5atrielle, 1*02 ( /rder XVosterogrammida Yilson, 200F (1hilognatha in"ertae sedis)&F2' Dn%ra"lass -entaKonia Brandt, 1*GG ( /rder X7mynilyspedida Ho%%man, 1I9I !uperorder 5ima"omorpha -o"o"k, 1*I$ ( /rder Glomeridesmida 1ook, 1*IF !uperorder /nis"omorpha -o"o"k, 1**E ( /rder Glomerida Brandt, 1*GG ( /rder !phaerotheriida Brandt, 1*GG Dn%ra"lass Helminthomorpha -o"o"k, 1**E !uperorder X7r"hipolypoda !"udder, 1**2 ( /rder X7r"hidesmida Yilson W 7nderson 200$ ( /rder X1o:iedesmida Yilson W 7nderson 200$ ( /rder X;uphoberiida Ho%%man, 1I9I ( /rder X-alaeosomatida Hannibal W 3rKeminski, 200F ( /rder X-leurojulida !"hneider W Yerneburg, 1II* (possibly sister to 1olobognatha)&9' !ubter"lass 1olobognatha Brandt, 1*G$ ( /rder -latydesmida 1ook, 1*IF ( /rder -olyKoniida 1ook, 1*IF ( /rder !iphono"ryptida 1ook, 1*IF ( /rder !iphonophorida 4e:port, 1*$$ !ubter"lass ;ugnatha 7ttems, 1*I* !uperorder 2uli%ormia 7ttems, 1I29 ( /rder 2ulida Brandt, 1*GG ( /rder !pirobolida 1ook, 1*IF ( /rder !pirostreptida Brandt, 1*GG ( !uper%amily XTyloiuloidea 1ook, 1*IF (!ometimes aligned :ith !pirobolida)&FG' !uperorder 4ematophora Qerhoe%%, 1I1G ( /rder 1allipodida -o"o"k, 1*I$

( ( ( !uperorder ,ero"haeta 1ook, 1*IF ( !ee also&edit' 7rthropods portal ( 5ea% litter ( !oil biology 6e%eren"es&edit'

/rder 1hordeumatida -o"o"k 1*I$ /rder !temmiulida 1ook, 1*IF /rder !iphoniulida 1ook, 1*IF /rder -olydesmida -o"o"k, 1**E

1 2ump upZ <,ost leggy millipede redis"o8ered<. BB1 4e:s. 7ugust *, 2009. 2 Z 2ump up to)a b " d !hear, Y. (2011). <1lass 0iplopoda de Blain8ille in Ger8ais, 1*$$. Dn) Vhang, V. [. (;d.) 7nimal biodi8ersity) 7n outline o% higher le8el "lassi%i"ation and sur8ey o% taxonomi" ri"hness<. VootaxaG1$*) 1FI+19$. G 2ump upZ <Possil millipede %ound to be oldest land "reature<. 144 (%rom6euters). 2E 2anuary 200$. $ 2ump upZ ,. G. 5o"kley W 1hristian ,eyer (201G). <#he tradition o% tra"king dinosaurs in ;urope<. 0inosaur #ra"ks and /ther Possil Pootprints o% ;urope. 1olumbia Jni8ersity -ress. pp. 2F+F2. D!B4 IE*02G1F0$90E. F Z 2ump up to)a b ,inelli, 7lessandroS !ergei D. Golo8at"h (2001). <,yriapods<. Dn !imon 7. 5e8in. ;n"y"lopedia o% Biodi8ersity. pp. 2I1+G0G.D!B4 012229*9F2. 9 Z 2ump up to)a b " d e % g h i j k l m !ier:ald, -etraS Bond, 2ason ;. (200E). <1urrent !tatus o% the ,yriapod 1lass 0iplopoda (,illipedes)) #axonomi" 0i8ersity and -hylogeny<. 7nnual 6e8ie: o% ;ntomology F2 (1)) $01+ $20.doi)10.11$9Rannure8.ento.F2.111*0F.0I0210. -,D0 1E19G*00. E 2ump upZ 5e:is, 2. G. ;. (200*). #he Biology o% 1entipedes (0igitally printed 1st paperba"k 8ersion. ed.). 1ambridge)1ambridge Jni8ersity -ress. pp. 110+111. D!B4 IE*0F210G$111. * Z 2ump up to)a b " d e % g 6obert 0. Barnes (1I*2). Dn8ertebrate Voology. -hiladelphia, -7) Holt !aunders Dnternational. pp. *1*+*2F. D!B4 0 0G 0F9E$E F. I 2ump upZ 2ohn 5. 1apinera, ed. (200*).<,illipedes<. ;n"y"lopedia o% ;ntomology. !pringer. pp. 2IGF+2GIE. D!B4 IE* 1 $020 92$2 1. 10 2ump upZ ,esibo8, 6obert. <-aranota<.;xternal 7natomy o% -olydesmida. 6etrie8ed G0 /"tober 201G. 11 Z 2ump up to)a b " d e !helley, 6o:land ,. (1III). <1entipedes and ,illipedes :ith ;mphasis on 4orth 7meri"an Pauna<. #he 3ansas !"hool

4aturalist $F (G)) 1+19. 12 Z 2ump up to)a b Blo:er, 2. Gordon (1I*F).,illipedes) 3eys and 4otes %or the Ddenti%i"ation o% the !pe"ies. 5ondon) -ublished %or the 5innean !o"iety o% 5ondon and the ;stuarine and Bra"kish Yater !"ien"es 7sso"iation by ;.2. Brill.D!B4 I00$0E9I*0. 1G 2ump upZ #amaris G. -inheiro, ,arin\K D. ,arOues W 5eandro 0. Battirola (200I). <5i%e "y"le o% -oratia sal8ator(0iplopoda) -olydesmida) -yrgodesmidae)<. Voologia (1uritiba)29 ($)) 9F*+992. doi)10.1FI0R!1I*$ $9E0200I000$00010. 1$ 2ump upZ ,esibo8, 6obert. <Gonopods<.;xternal 7natomy o% -olydesmida. 6etrie8ed 2E /"tober 201G. 1F 2ump upZ ;ngho%%, HenrikS 7kkari, 4esrine (2011). <7 "allipodidan "o"oon (0iplopoda, 1allipodida, !"hiKopetalidae)<. Dnternational 2ournal o% ,yriapodology F) $I+FG.doi)10.G*IERijm.F.1IIF. 19 2ump upZ 7dis, 2oa"him (1I*9). <7n <aOuati"< millipede %rom a 1entral 7maKonian inundation %orest<. /e"ologia 9* (G)) G$E+ G$I.doi)10.100ERBP010G9EGE. 1E 2ump upZ Burro:s, P.2.S 0. P. Hales, 7. 2. Beattie (1II$). <7Ouati" millipedes in 7ustralia) a biologi"al enigma and a "onser8ation saga<. 7ustralian Voologist 2I (G $)) 21G+219. 1* 2ump upZ Barber, 7.0. (;d) (201G). <Yorld 0atabase o% 5ittoral ,yriapoda<.Yorld 6egister o% ,arine !pe"ies. 6etrie8ed 2F /"tober 201G. 1I 2ump upZ Barker, G.,. (200$). <,illipedes (0iplopoda) and 1entipedes (1hilopoda)(,yriapoda) as predators o% terrestrial gastropods<. Dn G.,. Barker (ed.). 4atural enemies o% terrestrial mollus"s. Yalling%ord, /x%ordshire, J3) 17B Dnternational. pp. $0F+$29. D!B4 IE*0*F1II091G. 20 2ump upZ Yeldon, -aul 2.S 1ranmore, 1atherine P.S 1hat%ield, 2eni%er 7. (2009). <-rey rolling beha8ior o% "oatis (4asua spp.) is eli"ited by benKoOuinones %rom millipedes<.4atur:issens"ha%ten IG (1)) 1$+19.doi)10.100ERs0011$ 00F 009$ K. 21 2ump upZ !aporito, 6. 7.S 0onnelly, ,. 7.S Ho%%man, 6. 5.S Garra%%o, H. ,.S 0aly, 2. Y. (200G). <7 !iphonotid ,illipede (6hinotus) as the !our"e o% !piropyrroliKidine /ximes o% 0endrobatid Progs<. 2ournal o% 1hemi"al ;"ology 2I (12)) 2E*1+2E*9.doi)10.102GRB)2/;1.000000*09F.2*G9$.a0. 22 2ump upZ ;isner, #S ;isner, ,S 7ttygalle, 7BS 0eyrup, ,S ,ein:ald, 2 (1II*).<6endering the inedible edible) "ir"um8ention o% a millipedeLs "hemi"al de%ense by a preda"eous beetle lar8a.<. -ro"eedings o% the 4ational 7"ademy o% !"ien"es o% the Jnited !tates o% 7meri"a IF (G)) 110*+1G. -,D0 I$$*2IG. 2G 2ump upZ Dto, P. (1II*). <1olony "omposition and spe"ialiKed predati #emporal range) $2*+0,a -re. .

/ ! 0 1 - # 2 3 -g 4 5ate !ilurian to 6e"ent 6usty millipede (#rigoniulus "orallinus) !"ienti%i" "lassi%i"ation 3ingdom 7nimalia ) -hylum) 7rthropoda !ubphylu ,yriapoda m) 1lass) 0iplopoda 0e Blain8ille in Ger8ais, 1*$$ !ub"lasses ( -eni"illata ( 1hilognatha ( 7rthropleuridea (extin"t) 0i8ersity 19 orders, ". 12,000 spe"ies ,illipedes ("lass 0iplopoda) are myriapodous arthropods that ha8e t:o pairs o% legson most body segments. ;a"h double legged segment is a result o% t:o single segments %used together as one (the name <0iplopoda< "omes %rom the Greek :ords=>?@AB (diplous), <double< and ?A=CB (podos), <%oot<). ,ost millipedes ha8e 8ery elongated "ylindri"al or %lattened bodies :ith more than 20 segments, :hile pill millipedes are shorter and "an roll into a ball, like a pillbug. #he name <millipede< is a "ompound :ord %ormed %rom the 5atin roots mille(<thousand<) and pes (<%oot<). 0espite their name, no kno:n millipede has 1,000 legs, although the rare spe"ies Dlla"me plenipes has up to EF0.&1' 1ommon spe"ies ha8e bet:een G9 and $00 legs. #here are approximately 12,000 named spe"ies in "a. 1$0 %amilies.&2' #he longest spe"ies is the giant 7%ri"an millipede (7r"hispirostreptus

gigas). ,ost millipedes are slo: mo8ing detriti8ores, eating de"aying lea8es and other deadplant matter. Ho:e8er, they "an also be minor garden pests, espe"ially in greenhouses:here they "an "ause se8ere damage to emergent seedlings. ,illipedes "an be easily distinguished %rom the some:hat similar and related"entipedes (1lass 1hilopoda) :hi"h mo8e rapidly, are "arni8orous, and ha8e a single pair o% legs %or ea"h body segment. #he s"ienti%i" study o% millipedes is kno:n asdiplopodology, and a s"ientist :ho studies them is "alled a diplopodologist. 1ontents &hide' ( 1 ;8olution 2 1hara"teristi"s ( 2.1 Head ( 2.2 Body ( 2.G Dnternal organs ( G 6eprodu"tion and gro:th $ ;"ology ( $.1 Habitat ( $.2 0iet ( $.G -redators and parasites ( $.$ 0e%en"e me"hanisms ( F ,illipedes in relation to people ( 9 1lassi%i"ation ( E !ee also ( * 6e%eren"es ( I ;xternal links ;8olution&edit' #his "lass o% arthropod is thought to be among the %irst animals to ha8e "olonised land during the !ilurian geologi" period. #hese early %orms probably ate mosses and primiti8e 8as"ular plants. #he oldest kno:n land "reature, -neumodesmus ne:mani, :as a 1 "m (0.GI in) long millipede, and li8ed $2* million years ago.&G' Dn the Jpper 1arboni%erous (G$0 to 2*0 million years ago),7rthropleura be"ame the largest kno:n land in8ertebrate o% all time, rea"hing lengths o% up to 2.9 m (* %t 9 in). ,illipedes, "entipedes, and other terrestrial arthropods attained 8ery large siKes in "omparison to modern spe"ies in the oxygen ri"h en8ironments o% the 0e8onian and 1arboni%erous periods, and some "ould gro: larger than one metre. 7s oxygen le8els lo:ered through time, arthropods be"ame smaller in siKe.&$' 1hara"teristi"s&edit'

 #he 4orth 7meri"an millipede 4ar"eus ameri"anus + head :ith eyes, antennae, and enlarged "ollum ,illipede anterior anatomy ,illipedes range %rom 2 mm (0.0EI in) to around GF "m (1$ in) in length,&F' and "an ha8e as %e: as ele8en to o8er one hundred segments. #hey are generally bla"k or bro:n in "olour, although there are a %e: brightly "oloured spe"ies. #he millipedeLs most ob8ious %eature is its large number o% legs. Ha8ing many short legs makes millipedes rather slo:, but the many legs pushing in unison pro8ides po:er%ul strength %or burro:ing. Head&edit' #he head o% a millipede is typi"ally rounded abo8e and %lattened belo: and bears a pair o% large mandibles in %ront o% a plate like stru"ture "alled a gnatho"hilarium (<ja: lip<).&9' #he head "ontains a single pair o% antennae :ith se8en or eight segments and a group o% sensory "ones at the tip.&9' ,any orders also possess a pair o% sensory organs kno:n as the #MmMs8Nry organs, shaped as small o8al rings posterior and lateral to the base o% the antennae. #heir true %un"tion is unkno:n,&9' but they also o""ur in some "entipedes, and are possibly used to measure humidity or light le8els in the surrounding en8ironment. &E' ,illipede eyes "onsist o% a number o% simple %lat lensed o"elliarranged in a group on the side o% the head. ,any spe"ies o% millipedes, in"luding the entire order -olydesmida and "a8e d:elling millipedes su"h as 1auseyella and #ri"hopetalum, ha8e se"ondarily lost their eyes and are "ompletely blind.&F' Body&edit' -aranota o% a -olydesmidan (le%t) and-latydesmidan millipede #he body is %lattened or "ylindri"al, "omposed o% numerousmetemeri" segments, ea"h :ith an exoskeleton "onsisting o% a single "hitinous plate abo8e, one at ea"h side, and t:o or three on the underside. Dn many millipedes, these plates are %used to 8arying degrees, sometimes %orming a single "ylindri"al ring. #he plates are typi"ally hard, being impregnated :ith "al"ium salts.&*' Be"ause they la"k a :axy "uti"le, millipedes are sus"eptible to :ater

loss and must spend most o% their time in moist or humid en8ironments.&I' #he %irst segment behind the head is legless and kno:n as a "ollum. #he se"ond to %ourth body rings bear a single pair o% legs ea"h and are kno:n as <haplosegments< (%rom the Greek haplo) single). #he remaining segments, %rom the %i%th to the posterior, are properly kno:n as diplosegments, or double segments. ;a"h diplosegment bears t:o pairs o% legs, rather than just one as in "entipedes. #his is be"ause ea"h diplosgment is %ormed by the %usion o% t:o embryoni" segments. Dn some millipedes the last %e: segments may be legless. #he terms <segment< or <body ring< are o%ten used inter"hangeably to re%er to both haplo and diplosegments. #he %inal segment bears a telson, :hi"h "onsists o% a legless preanal ring, pair o% anal 8al8es ("loseable plates around the anus), and a subanal s"ale.&9'&*' ,illipedes in se8eral orders ha8e keel like extensions o% the body :all kno:n as paranota, :hi"h "an 8ary :idely in shape and siKe. #hey may allo: millipedes to :edge more se"urely into "re8i"es, prote"t the legs, or make the millipede more di%%i"ult %or predators to s:allo:.&10' Dnternal organs&edit' ,illipedes breathe through t:o pairs o% spira"les lo"ated 8entrally on ea"h segment.&11' ;a"h opens into an internal pou"h, and "onne"ts to a system o% tra"heae. #he heart runs the entire length o% the body, :ith an aorta stret"hing into the head. #he ex"retory organs are t:o pairs o% malpighian tubules, lo"ated near the mid part o% the gut. #he digesti8e tra"t is a simple tube :ith t:o pairs o% sali8ary glands to help digest the %ood. &*' 6eprodu"tion and gro:th&edit' ;pibolus pul"hripes mating ,illipedes sho: a di8ersity o% mating styles and stru"tures. Dn the basal order-olyxenida, males deposit spermatophores that are subseOuently pi"ked up by %emales.&11' Dn all other millipede groups, males possess one or t:o pairs o% modi%ied legs "alled gonopods :hi"h are used to trans%er sperm to the %emale during "opulation. #he lo"ation o% the gonopods di%%ers bet:een groups) in males o% the-entaKonia they are lo"ated at the rear o% the body and kno:n as telopods, :hile in Helminthomorpha + the 8ast majority o% spe"ies + they are lo"ated on the Eth body segment.&9' 7 %e: spe"ies are parthenogeneti", ha8ing %e:, i% any, males.&12'&1G' Gonopod morphology o% /xidus gra"ilis(-aradoxosomatidae). Palse "olor !;,image, s"ale bar) 0.2 mm Gro:th stages o% 4emasoma(4emasomatidae), :hi"h rea"hes reprodu"ti8e maturity in stage Q. #he mature gonopods de8elop %rom the legs o% segment E.

Gonopod morphology is the predominant means o% determining spe"ies among millipedes) the "omplex stru"tures di%%er mu"h bet:een spe"ies but 8ery little :ithin a spe"ies.&1$' #he genital openings are lo"ated on the third segment, and are a""ompanied in the male by one or t:o penes :hi"h deposit the sperm pa"kets onto the gonopods. Dn the %emale, the genital pores open into paired small sa"s "alled "yphopods or 8ul8ae, :hi"h are "o8ered by a small hood like "o8er, and are used to store the sperm a%ter "opulation.&*' ,illipede sperm is a%lagellate (la"ks a %lagellum), a uniOue trait among myriapods.&9' Pemales lay bet:een ten and three hundred eggs at a time, depending on spe"ies, %ertilising them :ith the stored sperm as they do so. ,any spe"ies simply deposit the eggs on moist soil or organi" detritus, but some "onstru"t nests lined :ith dried%ae"es, and may prote"t the eggs :ithin silk "o"oons.&*' Dn most spe"ies the %emale abandon the eggs a%ter laying but some spe"ies in the orders -latydesmida and!temmiulida pro8ide parental "are %or eggs and young.&11' #he young hat"h a%ter a %e: :eeks, and typi"ally ha8e only three pairs o% legs, %ollo:ed by up to %our legless segments. 7s they gro:, they "ontinually moult, adding %urther segments and legs as they do so. !ome spe"ies moult :ithin spe"ially prepared "hambers o% soil or silk,&1F' :hi"h they may also use to :ait out dry :eather, and most spe"ies eat the shed exoskeleton a%ter moulting. ,illipedes li8e %rom one to ten years, depending on spe"ies.&*' ;"ology&edit' Habitat&edit' Dn temperate Kones, millipedes are most abundant in moist de"iduous %orests, but they also o""ur in "oni%erous %orests, deserts, "a8es, and alpine e"osystems.&11' !ome spe"ies "an sur8i8e %resh:ater %loods and li8e submerged under:ater %or up to 11 months.&19' &1E' 7 %e: spe"ies o""ur near the seashore and "an sur8i8e in some:hat salty "onditions. &12'&1*' 0iet&edit' ,ost millipedes are detriti8ores and %eed on de"omposing 8egetation, %ae"es, or organi" matter mixed :ith soil. ,illipedes in the order -olyxenida graKe algae %rom bark, and -latydesmida %eed on %ungi.&9' 7 %e: spe"ies are omni8orous or o""asionally "arni8orous, %eeding on inse"ts, "entipedes, earth:orms, or snails&*'&1I' !ome spe"ies ha8e pier"ing mouth parts that allo: them to %eed on plant jui"es.&11' -redators and parasites&edit' ,illipedes are preyed upon by a :ide range o% animals, in"luding 8arious reptiles, amphibians, birds, mammals, and inse"ts.&9',ammalian predators su"h as "oatis and meerkats roll "aptured millipedes on the ground to deplete de%ensi8e se"retions and rub them o%% the body be%ore "onsuming,&20' :hile "ertain poison dart %rogs are belie8ed to in"orporate the toxi" "ompounds o% millipedes into their o:n de%enses.&21' !e8eral in8ertebrates ha8e spe"ialiKed beha8iors or stru"tures to %eed on millipedes, in"luding lar8al glo::orm beetles,&22' -robolomyrmex ants,&2G' "hlamydephorid slugs,&2$'

and the preda"eous dung beetle0elto"hilum 8algum.&2F' 4ar"eus ameri"anus "urled in de%ensi8e position -arasites o% millipedes in"lude nematodes, phaeomyiid %lies, and a"antho"ephalans. &9' 0e%en"e me"hanisms&edit' 0ue to their la"k o% speed and their inability to bite or sting, millipedesL primary de%en"e me"hanism is to "url into a tight "oil + prote"ting their deli"ate legs inside an armoured body exterior. ,any spe"ies also emit 8arious poisonous liOuid se"retions through mi"ros"opi" oKopores (also "alled odori%erous or repugantorial glands), along the sides o% their bodies as a se"ondary de%en"e. #hese se"retions may in"ludealkaloids, benKoOuinones, phenols, terpenoids, andRor hydrogen "yanide, among many others. &29'&2E' !ome o% these substan"es are "austi" and "an burn the exoskeleton o%ants and other inse"t predators, and the skin and eyes o% larger predators. -rimates su"h as "apu"hin monkeys and lemurs ha8e been obser8ed intentionally irritating millipedes in order to rub the "hemi"als on themsel8es to repel mosOuitoes.&2*'&2I'&G0'!ome o% these de%ensi8e "ompounds also sho: anti%ungal a"ti8ity.&G1' #he bristly millipedes (order -olyxenida) la"k both an armoured exoskeleton and odi%erous glands, and instead are "o8ered in numerous bristles that in at least one spe"ies, -olyxenus %as"i"ulatus, deta"h and entangle ants.&G2' ,illipedes in relation to people&edit' ,illipedes do not bite, and their de%ensi8e se"retions are mostly harmless to humans + usually "ausing only minor dis"oloration on the skin + but the se"retions o% some tropi"al spe"ies may "ause pain, it"hing, lo"al erythema, edema, blisters, e"Kema, and o""asionally "ra"ked skin.&GG'&G$'&GF'&G9' ;ye exposures to these se"retions "auses general eye irritation and potentially more se8ere e%%e"ts su"h as "onjun"ti8itis and keratitis.&GE' Pirst aid "onsists o% %lushing the area thoroughly :ith :aterS %urther treatment is aimed at relie8ing the lo"al e%%e"ts. 7 s:arm o% millipedes !ome millipedes are "onsidered household pests, in"luding Tenobolus "arni%ex :hi"h in%ests that"hed roo%s in Dndia.&G*' /ther spe"ies exhibit periodi"al s:arming beha8iour, :hi"h "an result in home in8asions,&GI' "rop damage,&$0' train delays, or e8en train "rashes and derailments.&$1'&$2' !ome o% the larger millipedes in the orders !pirobolida, !pirostreptida, !phaerotheriidaare popular as pets.&$G' !ome spe"ies "ommonly sold or kept in"lude spe"ies o%7r"hispirostreptus, 7phistogoniulus, 4ar"eus, and /rthoporus.&$$' ,illipedes also appear in %olklore and traditional medi"ine around the :orld. ,any

"ultures as"ribe millipede a"ti8ity :ith "oming rains.&$F' Dn the Uoruba "ulture o%4igeria, millipedes are used in pregnan"y and business rituals, and "rushed millipedes are used to treat %e8er, :hitlo:, and "on8ulsion in "hildren.&$9' Dn Vambia, smashed millipede pulp is used to treat :ounds, and in the Ba%ia people o% 1ameroon millipede jui"e is used to treat eara"hes.&$F' Dn "ertain Himalayan Bhotiya tribes, dry millipede smoke is used to treat hemorrhoids.&$E' 4ati8e people in ,alyasia use millipede se"retions in poison tipped arro:s.&$F' #he only reported usage o% millipedes as %ood by humans "omes %rom the Bobo people o%Burkina Paso, :ho "onsume boiled, dried millipedes in tomato sau"e.&$*' 1lassi%i"ation&edit' Por more details on this topi", see 5ist o% millipede %amilies. 7pproximate relati8e di8ersity o% extantmillipede orders, ranging %rom "a. G,F00 spe"ies o% -olydesmida to 2 spe"ies o%!iphoniulida. &2' 7pproximately 12,000 spe"ies o% millipede ha8e been des"ribed, but the true number o% spe"ies on earth has been estimated at up to *0,000.&9' ,illipedes are "onsidered a mega di8erse yet understudied group, :ith in"onsistent taxonomi" e%%ort o8er time.&$I' -olyxenus lagurus, a spe"ies o% the order -olyxenida Glomeris marginata, a pill millipede o% the order Glomerida /"toglena sierra, a spe"ies o% the order-olyKoniida 7nadenobolus monili"ornis, a spe"ies o% the order !pirobolida

 Harpaphe haydeniana, a spe"ies o% the order -olydesmida #he li8ing members o% the 0iplopoda are di8ided into sixteen orders in t:o sub"lasses.&2' #he basal sub"lass-eni"illata "ontains small spe"ies :hose exoskeleton is not "al"i%ied, and :hi"h are "o8ered in setae or bristles. 7ll other millipedes belong to the sub"lass1hilognatha, "onsisting o% t:o in%ra"lasses) the in%ra"lass -entaKonia"ontaining the short bodied pill millipedes, and the in%ra"lass Helminthomorpha (:orm like millipedes) "ontaining the great majority o% the spe"ies.&F0'&F1' #he higher le8el "lassi%i"ation o% millipedes is presented belo:, based on !hear, 2011,&2' and !hear W ;dge"ombe, 2010&F2' (extin"t groups). 6e"ent "ladisti" and mole"ular studies ha8e "hallenged traditional "lassi%i"ation s"hemes, and in parti"ular the position o% the orders !iphoniulida and -olyKoniida is not yet :ell established.&9'#he pla"ement and positions o% extin"t groups (X) kno:n only %rom %ossils is tentati8e and not %ully resol8ed.&9'&F2' 1lass 0iplopoda de Blain8ille in Ger8ais, 1*$$ !ub"lass -eni"illata 5atrielle, 1*G1 ( /rder -olyxenida Qerhoe%%, 1IG$ !ub"lass X7rthropleuridea (pla"ed in -eni"illata by some authors)&F2' ( /rder X7rthropleurida Yaterlot, 1IG$ ( /rder X;oarthropleurida !hear W !elden, 1IIF ( /rder X,i"rode"empli"ida !ub"lass 1hilognatha 5atrielle, 1*02 ( /rder XVosterogrammida Yilson, 200F (1hilognatha in"ertae sedis)&F2' Dn%ra"lass -entaKonia Brandt, 1*GG ( /rder X7mynilyspedida Ho%%man, 1I9I !uperorder 5ima"omorpha -o"o"k, 1*I$ ( /rder Glomeridesmida 1ook, 1*IF !uperorder /nis"omorpha -o"o"k, 1**E ( /rder Glomerida Brandt, 1*GG ( /rder !phaerotheriida Brandt, 1*GG Dn%ra"lass Helminthomorpha -o"o"k, 1**E !uperorder X7r"hipolypoda !"udder, 1**2

( /rder X7r"hidesmida Yilson W 7nderson 200$ ( /rder X1o:iedesmida Yilson W 7nderson 200$ ( /rder X;uphoberiida Ho%%man, 1I9I ( /rder X-alaeosomatida Hannibal W 3rKeminski, 200F ( /rder X-leurojulida !"hneider W Yerneburg, 1II* (possibly sister to 1olobognatha)&9' !ubter"lass 1olobognatha Brandt, 1*G$ ( /rder -latydesmida 1ook, 1*IF ( /rder -olyKoniida 1ook, 1*IF ( /rder !iphono"ryptida 1ook, 1*IF ( /rder !iphonophorida 4e:port, 1*$$ !ubter"lass ;ugnatha 7ttems, 1*I* !uperorder 2uli%ormia 7ttems, 1I29 ( /rder 2ulida Brandt, 1*GG ( /rder !pirobolida 1ook, 1*IF ( /rder !pirostreptida Brandt, 1*GG ( !uper%amily XTyloiuloidea 1ook, 1*IF (!ometimes aligned :ith !pirobolida)&FG' !uperorder 4ematophora Qerhoe%%, 1I1G ( /rder 1allipodida -o"o"k, 1*I$ ( /rder 1hordeumatida -o"o"k 1*I$ ( /rder !temmiulida 1ook, 1*IF ( /rder !iphoniulida 1ook, 1*IF !uperorder ,ero"haeta 1ook, 1*IF ( /rder -olydesmida -o"o"k, 1**E !ee also&edit' 7rthropods portal ( 5ea% litter ( !oil biology 6e%eren"es&edit' 1 2ump upZ <,ost leggy millipede redis"o8ered<. BB1 4e:s. 7ugust *, 2009. 2 Z 2ump up to)a b " d !hear, Y. (2011). <1lass 0iplopoda de Blain8ille in Ger8ais, 1*$$. Dn) Vhang, V. [. (;d.) 7nimal biodi8ersity) 7n outline o% higher le8el "lassi%i"ation and sur8ey o% taxonomi" ri"hness<. VootaxaG1$*) 1FI+19$. G 2ump upZ <Possil millipede %ound to be oldest land "reature<. 144 (%rom6euters). 2E 2anuary 200$. $ 2ump upZ ,. G. 5o"kley W 1hristian ,eyer (201G). <#he tradition o% tra"king

dinosaurs in ;urope<. 0inosaur #ra"ks and /ther Possil Pootprints o% ;urope. 1olumbia Jni8ersity -ress. pp. 2F+F2. D!B4 IE*02G1F0$90E. F Z 2ump up to)a b ,inelli, 7lessandroS !ergei D. Golo8at"h (2001). <,yriapods<. Dn !imon 7. 5e8in. ;n"y"lopedia o% Biodi8ersity. pp. 2I1+G0G.D!B4 012229*9F2. 9 Z 2ump up to)a b " d e % g h i j k l m !ier:ald, -etraS Bond, 2ason ;. (200E). <1urrent !tatus o% the ,yriapod 1lass 0iplopoda (,illipedes)) #axonomi" 0i8ersity and -hylogeny<. 7nnual 6e8ie: o% ;ntomology F2 (1)) $01+ $20.doi)10.11$9Rannure8.ento.F2.111*0F.0I0210. -,D0 1E19G*00. E 2ump upZ 5e:is, 2. G. ;. (200*). #he Biology o% 1entipedes (0igitally printed 1st paperba"k 8ersion. ed.). 1ambridge)1ambridge Jni8ersity -ress. pp. 110+111. D!B4 IE*0F210G$111. * Z 2ump up to)a b " d e % g 6obert 0. Barnes (1I*2). Dn8ertebrate Voology. -hiladelphia, -7) Holt !aunders Dnternational. pp. *1*+*2F. D!B4 0 0G 0F9E$E F. I 2ump upZ 2ohn 5. 1apinera, ed. (200*).<,illipedes<. ;n"y"lopedia o% ;ntomology. !pringer. pp. 2IGF+2GIE. D!B4 IE* 1 $020 92$2 1. 10 2ump upZ ,esibo8, 6obert. <-aranota<.;xternal 7natomy o% -olydesmida. 6etrie8ed G0 /"tober 201G. 11 Z 2ump up to)a b " d e !helley, 6o:land ,. (1III). <1entipedes and ,illipedes :ith ;mphasis on 4orth 7meri"an Pauna<. #he 3ansas !"hool 4aturalist $F (G)) 1+19. 12 Z 2ump up to)a b Blo:er, 2. Gordon (1I*F).,illipedes) 3eys and 4otes %or the Ddenti%i"ation o% the !pe"ies. 5ondon) -ublished %or the 5innean !o"iety o% 5ondon and the ;stuarine and Bra"kish Yater !"ien"es 7sso"iation by ;.2. Brill.D!B4 I00$0E9I*0. 1G 2ump upZ #amaris G. -inheiro, ,arin\K D. ,arOues W 5eandro 0. Battirola (200I). <5i%e "y"le o% -oratia sal8ator(0iplopoda) -olydesmida) -yrgodesmidae)<. Voologia (1uritiba)29 ($)) 9F*+992. doi)10.1FI0R!1I*$ $9E0200I000$00010. 1$ 2ump upZ ,esibo8, 6obert. <Gonopods<.;xternal 7natomy o% -olydesmida. 6etrie8ed 2E /"tober 201G. 1F 2ump upZ ;ngho%%, HenrikS 7kkari, 4esrine (2011). <7 "allipodidan "o"oon (0iplopoda, 1allipodida, !"hiKopetalidae)<. Dnternational 2ournal o% ,yriapodology F) $I+FG.doi)10.G*IERijm.F.1IIF. 19 2ump upZ 7dis, 2oa"him (1I*9). <7n <aOuati"< millipede %rom a 1entral 7maKonian inundation %orest<. /e"ologia 9* (G)) G$E+ G$I.doi)10.100ERBP010G9EGE. 1E 2ump upZ Burro:s, P.2.S 0. P. Hales, 7. 2. Beattie (1II$). <7Ouati" millipedes in 7ustralia) a biologi"al enigma and a "onser8ation saga<. 7ustralian Voologist 2I (G $)) 21G+219.

1* 2ump upZ Barber, 7.0. (;d) (201G). <Yorld 0atabase o% 5ittoral ,yriapoda<.Yorld 6egister o% ,arine !pe"ies. 6etrie8ed 2F /"tober 201G. 1I 2ump upZ Barker, G.,. (200$). <,illipedes (0iplopoda) and 1entipedes (1hilopoda)(,yriapoda) as predators o% terrestrial gastropods<. Dn G.,. Barker (ed.). 4atural enemies o% terrestrial mollus"s. Yalling%ord, /x%ordshire, J3) 17B Dnternational. pp. $0F+$29. D!B4 IE*0*F1II091G. 20 2ump upZ Yeldon, -aul 2.S 1ranmore, 1atherine P.S 1hat%ield, 2eni%er 7. (2009). <-rey rolling beha8ior o% "oatis (4asua spp.) is eli"ited by benKoOuinones %rom millipedes<.4atur:issens"ha%ten IG (1)) 1$+19.doi)10.100ERs0011$ 00F 009$ K. 21 2ump upZ !aporito, 6. 7.S 0onnelly, ,. 7.S Ho%%man, 6. 5.S Garra%%o, H. ,.S 0aly, 2. Y. (200G). <7 !iphonotid ,illipede (6hinotus) as the !our"e o% !piropyrroliKidine /ximes o% 0endrobatid Progs<. 2ournal o% 1hemi"al ;"ology 2I (12)) 2E*1+2E*9.doi)10.102GRB)2/;1.000000*09F.2*G9$.a0. 22 2ump upZ ;isner, #S ;isner, ,S 7ttygalle, 7BS 0eyrup, ,S ,ein:ald, 2 (1II*).<6endering the inedible edible) "ir"um8ention o% a millipedeLs "hemi"al de%ense by a preda"eous beetle lar8a.<. -ro"eedings o% the 4ational 7"ademy o% !"ien"es o% the Jnited !tates o% 7meri"a IF (G)) 110*+1G. -,D0 I$$*2IG. 2G 2ump upZ Dto, P. (1II*). <1olony "omposition and spe"ialiKed predati