commonname: rai nbowscar ab

scient i
fi
cname: Phanaeusv indexMacl eay(Insecta:Coleoptera:Scar abaeidae)
Intr
oduct ion-Sy nony my-Di stri
buti
on-Descr i
pti
on-Li feCy cle-Di et-I mpor tance-
Conser v
ation-Sel ectedRef er ences
Intr
oduct ion( Backt oTop)
Rainbow scar abs,Phanaeus v i
ndex MacLeay ,ar e member s oft he beet l
ef amily
Scar abaeidae,whi chal ongwi t
ht hefamilyGeot rupidae,arecommonl yknownasdung
beet l
es( Bertoneetal .2004) .Scar abbeetleswer etheobj ectsofwor shipi nAncientEgy pt
andt hemor espect acularv arieti
esaremadei ntojewelry(Woodr uff1973) .TheAf ri
can
dung- rol
li
ngbeet leut il
izesthesun, moonandst arsasgui desf orwal ki
ngi nastraightli
ne
(Dackeetal .2013) .ThegenusPhanaeusMacLeayi sdistr
ibutedpr i
mar i
lyinNeot ropical
habi t
atswi th100speci es,butal soi ntheNear cti
cr egi
onwi t
hni nespeciesandt heWest
Indieswi thone.Dungbeet l
esser veanimpor tantroleinpast ureecosy stems,whi chhas
resultedi nt heiri ntroducti
onar oundt hewor l
d.Ther ainbow scar ab,Phanaeusv i
ndex
MacLeay ,hasabr i
ghtex teriorofmet alli
cgr een,blue,andr edi nterspersedwi t
hgol den
refl
ect i
ons( Figures1and2) .

Fi
gur
e1.Red-
greenmal
ePhanaeusv
index.Phot
ogr
aphbyPaul
Skel
l
ey.

Synonymy( Backt
oTop)
Scarabaeuscarni
fexLi
nnaeus1767
Phanaeusv i
ndexMacleay ,
1819
Phanaeusv i
ndexcyanel
lusRobinson,1938
Phanaeusv i
ndexruber
virensRobinson,
1948
(Ar
naud2002, Edmonds1994)

Dist
ri
bution(
BacktoTop)
Therainbowscarabi
snati
vetoandf oundextensi
vel
yin,t
heeasternUnit
edStat
esfr
om
Massachusett
stoSouthDakotaint
henor t
handTex astoFlori
dainthesout
h(Woodruf
f
1973).TheFlori
daKeysandt heEvergl
adesar etheonlyregi
onsinFlori
dawherethe
rai
nbowscarabhasnotbeenfound(Woodruff1973).
Descr
ipt
ion(BacktoTop)
Lar
vae:Thel
arvaearegrubl
ike,
wit
hacharact
eri
sti
cC-shapedbody(Rat
cli
ff
eetal
.2002)
.
Theheadishardenedandthebodyissof
tandpl
iabl
e(Woodruff1973)
.

Adul t
s: Theadul tsarebul ky, withanov alorelongat eshape.Fr om thet opoft heheadt o
thebackoft heabdomen, mal esandf emal esar eabout1t o2. 2centimet erslong
(Bertoneetal .2004) .Theyhav elamel l
ateant ennae.Themal esar eeasi lyidentif
iedby
theirmet al l
i
cappear anceandal argehor nar i
singf r
om thedor salsurfaceoft hehead.A
similarspeci es, Phanaeusi gneus( MacLeay )
,isalsomet alli
c,buthasasmal l
erhor n.
Somemal esofPhanaeusv index, calledmi normal es,hav egr eatl
yreducedhor ns,butan
exami nationoft hedi f
ferencesi nt hepost eri
orabdomi nal segment scanbeusedi n
determi ningt hegender .Thef emal esdonotpossesshor ns.Thedor sal surfaceoft he
pronot um ( thor ax)ispl at
e- l
ike,wi t
ht hepost eri
orendsoft hemal e'spronot um pointed
andsl ightlyr aisedabov et heel ytra( modi fi
edf orewingsofbeet l
es,hardenedi nto
protectiveshi eldsfort hehi ndwi ngs) .Thef emal e'spronot um, bycont r
ast ,isnotr aised
abov et heel yt r
a.Thepr onot um andel ytr
ai nbot hthemal eandf emal ehav edistinctiv
e
coloration.Thepr onotum gener all
yhasamet alli
cbr onzeandr edcol oration.Theel y t
ra
gener all
yar eamet alli
cgr een.Ther ear erareindi v
idualsthatar edar kbl uet oblacki n
color,lackingal lbronz e-r
edcol orat i
on( Figures3and4)( Woodr uff1973) .

Fi
gur
e2.Red-
greenf
emal
ePhanaeusv
index.Phot
ogr
aphbyPaul
Skel
l
ey.
 Fi
gur
e3.Bl
ue-
blackmal
ePhanaeusv
index.Phot
ogr
aphbyPaul
Skel
l
ey.

Fi
gur
e4.Gr
een-
bluef
emal
ePhanaeusv
index.Phot
ogr
aphbyPaul
Skel
l
ey.

Lif
eCy cl
e( Backt oTop)
Basedonnestconst r
uctionbehav ior,therear et hreet ypesofdungbeet les:t unnelers
(paracoprids),dwel l
ers( endocopri
ds) ,andr oll
ers( tel
ecopr i
ds).Phanaeusv indexi san
exampl eofat unneler(Ratcl
iff
eetal .2002) .I
nt unnel i
ngspeci es,bot
hsex escr eatea
tubebyex cavat i
ngthesoi lunderneat hthedungpat ,culminati
ngi nachamberbel owt he
surfaceoft hedungpat( Fi
gure5) .Thet unnelingexcav ati
onsbr i
ngsubsur facesoi l
,
call
edsoi lcasts,tothegr ound'
ssur face.Thebr oodbal li
smadeunder neatht hedungpat
(Ratcli
ffeetal.2002) .Thebr oodbal lconsi stsofmoi stdungcoat edwit
hsoi ltoser veas
foodf orthelarv aandy oungadul t(Bertoneetal .2004) .Thebal lofdungi spul l
edintot he
tunnel,af ew cent i
met ersbelow gr ound,andf ormedi ntobrood,orf eedingbal ls.The
tunnelsar ef i
ll
edwi t
hl oosesoiltopr otectt hebr oodbal latthebot t
om oft het unnel
(Bertoneetal .2006) .Becauset unnel ersbr ingthedungunder ground,thet unnelersar e
themostsoughtaf terdungbeet l
esf ordungdegr adat i
on( KaufmanandWood2012) .

Thebr oodballi
sshapednonunif
ormly,t
houghitoftenappearspyrif
orm orpear
-shaped.
Priortomakingthebroodbal
l,Phanaeusvindexperfor
msa' butti
ng'act
iononthedung,
therebypushi
ngi tawayf
rom theori
ginaldeposi
ti
onsite(Half
fterandMat t
hews1966) .
Whent hef emal ebegi nst or olet hedung,t hisact ionat tractsamal e.Thenewl yf ormed
couplewor kt ocompl eteabal lofdungandal sot oconst ructabur row,whichmayser v
e
asanov ipositionsi teorsi mpl yasashel t
er( Half fteretal .1974,Hal ff
terandMat thews
1966,Thomas2001) .Whenf emal eov i
posit
ionoccur s,aneggi splacedatt hepoi ntof
thebroodbal l( Woodr uff1973) .Oneorsev eralbr oodbal lsarepl acedint hebur rows
whichconsi stofdungcoat edwi thear t
htopr eser vet hedungqual i
ty(Half
fteretal.1974) .
Inadditi
ont ot het unnelconst r
uct edf orthebr oodbal l(thepr i
mar ytunnel),asecond
tunneli
sconst ructedandst ockedwi t
hadungbal lf orfoodf orthenewl yemer gedadul t
s
andov er
wi nteringadul ts(thesecondar ytunnel )( Fincher1972) .Onceov i
positi
oni nt he
broodbal liscompl ete,thef emal eceasest ocar efort heegg( andlatert hel arva),
alt
hought hef emal eandhermat eremai ninthebur row( Halff
teretal.1974).
Afterhatching,al arvadev el
opswi thinthebr oodbal l(Fincher1972) .Mol ti
ngoccur s
twiceduringt hel arvalstage,af terwhi chitpupat es( transi t
ionstoadul tstages)wi thi
n
thebroodbal l.Inar easnor thofFl or i
da,overwi nteringl arvaer emainbelowt hef r
ostl i
ne
andr i
sefrom hi bernationuponf eelingtheincr easi ngt emper aturesofspr i
ng.Theadul t
s
ofPhanaeusv indexhav ealsobeenknownt oov erwi nter( BlumeandAga1976) .

Thef i
nalst agei nt hel i
fecy cl
ei st headul tstage,whi chst artsi nt hepr i
mar yand
secondar ytunnel s.Theadul temer gesfrom t hetunneli nr esponset oei thert emperature
orr ainfall(
Rat cli
ffeetal .2002) .Afteremer gence, adul t
swi lleventuall
ysear chforamat e.
Followi ngmat ing,femal est henr epeatt heegg- layingcy cle.Adul tsbur yandconsume
dungaspar toft heirdi et,therebypr ovi
dingv aluablenut r
ientr ecycl
ingi npast uresystems
(Ber toneetal .2006) .Soi lst ructureandcomposi ti
onseem t oplayar olei nPhanaeus
vindexsur vi
v alanddi st r
ibution.Sandysoi l
swer ef oundt obedel eterioust oPhanaeus
vindexbycausi nglar val desiccation( Bertoneetal .2006, Fincher1973) .
Int hel aboratory,thet otalt imef rom eggt oadul temer gencei st wot osi xmont hs
(Fincher1972) .Underl aborat orycondi t
ions,Phanaeusv indexadul t
scanl i
v eforovera
year .Underf ieldcondi ti
ons, sur vi
valismostl ikel
ymuchl esst hanay ear( Fincher1972) .
Onedocument edpr edat orofPhanaeusv i
ndexi st hebur rowi ngowl ,At henecuni culari
a
(Mol ina),whi ch has been f ound t o hav e Phanaeus v index f ragment si ni t
s dung
(Woodr uff1973) .

Thepr edator
ymi teMacr ochelesamy gdali
gera(Berlese)isof t
enf oundont headult
rai
nbowscar ab.Thispredat
orymiteeatsinsecteggs,smallinsectl
arvae,andnemat odes.
Macrochelesamy gdali
geraappearstout i
li
zethedispersalcapabil
it
iesoft hebeetl
eina
symbioti
crelati
onshipknownasphor esy,beingt
ransportedfrom dungpatt odungpatby
cli
ngi
ngt othebeetl
e( Woodruff1973).
 Fi
gur
e5.Li
fehi
stor
yofPhanaeusv
index
.Wat
ercol
orbyXav
ierMoss.

Diet( Backt oTop)

Dungbeet l
es,i ngener al,hav ebeenshownt oprefercer t
aint y
pesofani maldung,whi ch
arel istedi ndescendi ngor der :swine,opossum,dog,cow,r accoon,andhor se( Bertone
2004,Fi ncheretal .1970) .The swi ne and opossum dung was consi der ablymor e
attract i
vet han the ot hers( Woodr uff1973) .Phanaeus v i
ndex al so has a st r
ong
pref erence f orhuman dung ( Woodr uff1973) .Dung beet l
es,asa whol e,occupya
compl exsetofni chesi nt heecosy stem r el
atedt ot hel ocationsofdungi nv arious
habi tats,soi ltypes,and seasonsofact ivi
ty.A st udyi n Ger many( Wassmer1995)
i
ndi cat edt hatsomedungbeet lespecieswer emor elikelytobef oundint hef orestrather
thant hepast ure,inthepast ur erathert hant hef orest ,andsomewer ef oundi nbot h
forestandpast urehabi tat
s.Howev er,seasonal it
ywasf oundt obemuchmor ecriti
calfor
ex plainingdungbeet lespeci esdi versit
yt henhabi tatni ches( Wassmer1995) .Phanaeus
vindex’ swi dedi str
ibuti
oni sduet oi t
st olerancef orawi dev arietyofsoilty pes( Fincher
1973) .I n east ern Tex as,Phanaeus v indexappear st o hav e a preference f oropen
past urest hathav esandysoi ls(PriceandMay2009) .Thi siscur iousduet osandysoi l
bei ngdel eter i
ousf orPhanaeusv index '
ssur vival(Fi ncher1973) .InTexas,Phanaeus
difformi sLeCont eof t
encoex istswi t
hPhanaeusv index.Whent hi
soccur s,Phanaeus
vindex' st endencyt oinhabi tsandysoi lsisov errul
edandPhanaeusv indexi nhabi t
scl ay
soi l
s( BlumeandAl ga1976,1978) .I nFlor i
da,asi nTex as,aspeci escoex i
stswi th
Phanaeusv index:Phanaeusi gneusMacl eay .Phanaeusi gneust endst oi nhabi tsandy
soi l
s, whi l
ePhanaeusv indexi nhabitsclaysoi ls(Edmonds1994) .

I
mportance(
BacktoTop)
I
nfest
ati
onofhumanswi t
hdungbeet
lesisar
areoccur
rence.Adi
ff
erentdungbeet
leof
t
hegenusCopr i
sandf amil
yScar
abaei
daehasbeenreportedt
oinfesthumansinSri
Lanka( Rajapakse1981) .Themostl ikel
ymechani sm fori nfest ationofdungbeet lesis
i
ngest ionoft helar vae,whi lethechanceofanov ipositi
onev entt hr ought heanusi sl ow
(Rajapakse1981) .Ther ehav ebeeni ncident swher edungbeet lesacci dentall
yi nfested
humans,suchasoccur redwithagr oupof186boyscout sinPennsy l
vaniai n1957.The
beet l
escr awl edi ntot heear soft heboyscout swhi letheywer esl eeping.Bl eedingwas
i
nduced byt hehi nd t i
bialspinesoft hebeet l
es,butno secondar yi nfectionswer e
repor t
ed( Mat tuckandFehn1958) .
Phanaeus v i
ndex i s known t o be t he i ntermedi ate host of a st omach wor m,
Phy socerphal us sexal atus Mol i
n,a nemat ode t hati nfects swi ne.Swi ne obt aint he
stomach wor m af teri ngesti
ng adul tPhanaeusv i
ndex( Fincheretal .1969) .I nt he
l
abor atory,thepr edat or ymi t
eMacr ochel esamy gdali
ger a,feedsoneggsoft hest omach
wor m( Woodr uff1973) .
Dungbeet lesar ei mpor tantinreduci ngwast eaccumul ationsofv ertebrates,par t
icularl
y
rumi nants( cattl
e,sheep,goat s,etc.)
.Thi swasdr amat i
cal l
ydepi ctedi nAust rali
a,whi ch
priortot hecol oni zationoft hecont inentbyEur opeans,washomet omar supialsandno
rumi nants.Theaccumul at
edwast eofi ntroducedr uminant sgr adual lyresultedi nacr isi
s
thatwasal lev i
atedbyt heintroductionofsev eralspeciesofr umi nant-specializi
ngdung
beet l
est otheAust ral
iancont i
nent( Bor nemi ssza2006, Tyndal e-Biscoe1996) .

Likemanydungbeet les,Phanaeusv i
ndexwasf oundt oenri
chclaysoil
sasar esul
tof
feedingonadungpi l
e.Consumpt ionofdungbyadul tbeetl
esprovi
desvaluabl
enut r
ient
recycli
ngi
npasturesystems(Ber t
oneetal.2006).Fur
thermore,
harmfulf
lyspeciessuch
ast hehornf
ly,Haemat obi
airr
itans(Li
nnaeus),maybeout competedbypopulat i
onsof
dungbeetlel
arvae(Bert
one2004, Bert
oneetal.2004).

Conser v
ation( Backt oTop)
Try ingt opr otectcat tlefrom har mf ulparasit
es, whileatthesamet imemai ntainingbeet l
e
popul at i
ons,r equiresgr eatpr udence.Manypest i
cidesusedt ot
reatpar asiti
cwor msi n
l
ivest ockwi ll,ifusedf oral ongdur ationandwi thoutcar ef
ulmanagement ,affectdung
beet l
esadv ersel y(Floate1998,Lumar etandEr rouissi2002). Theor alf ormul ationsof
i
v ermect inhav ebeenshownt obel essdev astati
ngt onont ar
get s,whi chi si nst ar k
cont rastt ot hesust ai
ned-rel easebol usofi vermect in,whichhaspr olongedef f
ect son
nont ar gets( Lumar etandEr roui ssi 2002).
Sel ect edRef er ences( Backt oTop)
Ar naudP.2002.Lescol eopter esdumonde.Vol 28.Phanaei ni.Hi
l
lsideBooks.Cant erbur y.
151pp.
Ber toneMA.2004.DungBeet les( Coleoptera:Scar abaeidaeandGeot rupridae)ofNor th
Car ol inacat tlepast uresandt heirimpl i
cati
onsf orpast ur
ei mprov ement .M. S.Thesi s.
Nor t hCar oli
naSt ateUni versity.Ral eigh,NC.159pp.
Ber toneMA, Gr eenJT, Washbur nSP, PooreSH,andWat sonDW.2006.Thecont ribution
oft unnel ing dung beet l
es t o past ure soilnut ri
ti
on.Pl antManagementNet wor k
Inter nat ional: ForageandGr azingl ands.(31Oct ober2016)
Ber toneMA,Wat sonW,St ringham M,Gr eenJ,Washbur nS,Poor eM,andHucksM.
(2004) .Dung beet l
es ofcent r aland east ern Nor th Carol
ina cattle past ures.Nor th
Car olinaCooper at iv eEx tensi on.Nor thCar olinaSt at eUni v ersity .Ralei gh, NC.( 31Oct ober
2016)
Blume RB,Aga A.1976.Phanaeus di ff ormi s Lecont e( Col eopt era:Scar abaei dae):
Clar i
f i
cat i
onofpubl isheddescr ipt ions,not esonbi ology ,anddi stribut i
oni nTexas.The
Coleopt eristsBul letin30: 199- 205.
BlumeRR,AgaA.1978.Obser v ationsonecol ogi calandphy logenet icr elationshi psof
Phanaeusdi f
for mi sLeCont eandPhanaeusv indexMacLeay( Col eopet era: Scar abaei dae)
i
nNor thAmer ica.Sout hwest ernEnt omol ogi st3: 113- 120.
Bor nemi ssz aG.( 2006) .We' v edungi tagai n!Ourexot icsol utiont ot hedungpr oblem in
Aust r alia.CSI RO.( nol ongerav ai l
abl eonl i
ne)
DackeM, Bai rdE, By rneM, Schol tzCH, War rantEJ.2013.Dungbeet l
esuset heMi lkyWay
foror ient ation.Cur rentBi ology23: 1- 3.
EdmondsWD.1994.Rev i
sionofPhanaeusMacl eay ,aNewWor ldgenusofScar abaeine
dungbeet l
es( Col eopt era: Scar abaei dae, Scar abaei nae) .Cont ributionsi nSci ence, Nat ural
Hist or yMuseum ofLosAngel esCount y443: 1- 105.
FincherGT.1972.Not esont hebi ologyofPhanaeusv i
ndex( Col eopt era:Scar abaei dae).
Jour nal oft heGeor giaEnt omol ogi cal Societ y7: 128- 133.
FincherGT.1973.Ni difi
cat i
on and r eproduct i
on ofPhanaeusspp.i nt hr eet extural
classesofsoi l (Col eopt era: Scar abaei dae).TheCol eopt eristsBul letin27: 33- 37.
FincherGT, Stewar tTB, andDav i
stR.1969.Beet lei nter medi at ehost sf orswi nespi ruri
ds
i
nsout her nGeor gi a.TheJour nal ofPar asi tology55: 355- 358.
FincherGT,St ewar tTB,andDav isR.1970.At tract ionofcopr ophagousbeet l
est of eces
ofv ar iousani mal s.Jour nal ofPar asi t
ology56: 378- 383.
Float eKD.1998.Of f-t
ar getef f ect sofi ver mect inoni nsect sandondungdegr adat ionin
sout her nAl ber ta, Canada.Bul let i
nofEnt omol ogi cal Resear ch88: 25- 35.
Hal ffterG,Hal fft erV,andLopezI G.1974.Phanaeusbehav i
or :foodt r
anspor tationand
bisexual cooper at ion.Env ironment al Entomol ogy3: 341- 345.
Hal ffterG,Mat thewsEG.1966.Thenat ur alhi st or yofdungbeet lesoft hesubf ami ly
Scar abaei nae( Col eopt era: Scar abaei dae).Fol iaEnt omol ógi caMex icana12- 14: 1-313.
Kauf man PE,Wood LA.2012.I ndigenous and exot i
c dung beet les ( Col eopt era:
Scar abaei dae and Geot r
upi dae)col l
ected i nf l
or ida cat tle past ures.Annal s oft he
Entomol ogical Soci etyofAmer ica105: 225- 231.
Lumar etJP, Er roui ssiF.2002.Useofant hel mi nticsi nher bi vor esandev aluat i
onofr i
sks
fort henont ar getf aunaofpast ur es.Vet erinar yResear ch33: 547- 562.
Mat t uckDR, FehnCF.1958.Humaneari nv asionsbyadul tScar abaei dbeet les.Jour nalof
Economi cEnt omol ogy51: 546- 547.
Price DL, May M. 2009. Behav i
or al ecol ogy of Phanaeus dung beet l
es
(Col eopt era:Scar abaei dae) :Rev i
ew and new obser vat i
ons.Act a Zool ógi ca Mexi cana
(nuev aser ie)25: 211- 238.
RajapakseS.1981.Beet l
emar asmus.Br itishMedi cal Jour nal 283: 1316- 1317.
Rat cliffeBC,JamesonML,Smi thABT.2002.Fami ly34.Scar abaei daeLat reill
e1802.I n:
Arnet tRH Jr ,ThomasMC,Skel leyPE,Fr ankJH ( eds. )Amer i
canBeet les,Vol ume2:
Poly phaga: Scar abaeoi deat hroughCur culionoi dea.CRCPr ess, BocaRat on, 861pp.
ThomasML.( 2001) .Dungbeet lebenef itsi nt hepast ureecosy stem.ATTRA.( 31Oct ober
2016)
Tyndal e-
Bisoce M.1996.Aust r
alia'
si ntroduced dung beet les:Or i
ginalreleases and
di
stributi
ons.CSI ROEntomologyTechni cal Repor t
.No.62.149pp.
WassmerT.1995.Sel ecti
on oft hespat ialhabi t
atofcopr ophagousbeet l
esint he
KaiserstuhlareanearFrei
burg( SW- Germany ).ActaEcol ogica16: 461-478.
Woodr uffRE.1973.Ar thropodsofFl orida and nei ghbor ing land areas.TheScar ab
BeetlesofFl ori
da,Vol
ume8.Fl oridaDepar tmentofAgr i
cultureandConsumerSer vices.
Contributi
onNo.260, BureauofEnt omol ogy, Gainesvil
le.220pp.
Woodr uffRE.1973.Thescar abbeet l
esofFl ori
da.Par tI ,theLapr ostict
i(subfami l
i
es
Scarabaeinae,Aphodi i
nae,Hy bosorinae,Ochodaei nae,Geot rupinae,Acanthocerinae).
Fl
or i
daDepar tmentofAgr i
cult
ur eandConsumerSer v i
ces.