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Appendix(S1:(Systematics(Of(The(Micrurus'fulvius(Complex(And(Taxonomic( Revision(Of(Micrurus'tener(

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Appendix(S1:(Systematics(Of(The(Micrurus'fulvius(Complex(And(Taxonomic( Revision(Of(Micrurus'tener( Streicher et al. 1 Appendix(S1:(Systematics(of(the(Micrurus'fulvius(complex(and(taxonomic( revision(of(Micrurus'tener( Introduction(( Coralsnakes*of*the*genus*Micrurus*Wagler*1824*from*North*and*Central*America*have*a* complicated*taxonomic*history,*likely*because*they*have*a*highly*conserved*morphology* (Boulenger*1896;*Schmidt*1933;*1958;*Slowinski*1995)*and*many*species*possess*color* pattern*polymorphism*(Schmidt*1958;*Roze*1996;*Campbell*and*Lamar*2004).*Although* molecular*data*have*been*used*to*explore*enzyme*diversity*in*venoms*(e.g.*Tanaka*et*al.* 2010;*Renjifo*et*al.*2012;*Margres*et*al.*2013;*CarbajalSSaucedo*2013),*most* phylogenetic*analyses*of*DNA*for*species*involved*in*the*Micrurus'fulvius*(Linneaus* 1776)*complex*(sensu*Castoe*et*al.*2012)*have*been*restricted*to*the*nominate*form* (Slowinski*1995;*Castoe*et*al.*2007;*Pyron*et*al.*2011,*2013)*or*this*and*M.'tener*(Baird* and*Girard*1853).*Renjifo*et*al.*(2012)*found*M.'fulvius*and*M.'tener*forming*a* monophyletic*group*sister*to*M.'diastema*(Duméril,*Bibron,*and*Duméril*1854),*another* species*found*in*Mexico.*Thus,*the*relatedness*of*these*morphologically*similar*snakes* remains*uncertain*due*to*low*species*coverage*with*at*least*16*species*occurring*in* Mexico.** In*the*main*text*we*present*evidence*that*M.'tener*is*a*species*comprised*of* individuals*that*possess*one*of*two*divergent*mitochondrial*haplogroup*types,*but* collectively*have*nuclear*DNA*variation*consistent*with*a*single*species*that*recently* expanded*northward.*Although*it*is*beyond*the*scope*of*our*study*to*discuss*the* Streicher et al. 2 systematics*of*all*North*American*Micrurus*in*monographic*form*(which*we*believe*is* desperately*needed),*our*molecular*findings*have*important*implications*for*the* systematics*of*these*snakes.*Our*expanded*sampling*of*species*from*Mexico*warrants* discussion*given*that*the*distribution*of*genetic*variation*challenges*(1)*currently* hypothesized*relationships*and*distributions*and*(2)*the*validity*of*several*taxa.*We* provide*below*an*overview*of*the*taxonomic*implications*from*our*study*and* justifications*for*synonymizing*M.'bernadi'(Cope*1887)*and*M.'tamaulipensis*LavinS Murcio*and*Dixon*2004*with*M.'tener.*We*also*recommend*not*recognizing*the*multiple* subspecies*that*have*been*described*from*this*taxon.* Subspecies,(geographic(range(extent,(and(close(relatives(of(Micrurus'tener(( Our*sampling*included*representatives*of*all*subspecies*of*M.'tener'currently* recognized,*M.*t.*tener,*M.*t.'fitzingeri*(Jan,*1858),*M.'t.*maculatus*Roze*1967,*and*M.*t.* microgalbineus*Brown*and*Smith*1942*(Table*S1).*We*sampled*60*individuals*of*M.'t.' tener,*three*individuals*of*M.'t.'fitzingeri*(M51,*M432,*and*M516),*two*individuals*of*M.' t.*maculatus*(M206*and*M332;*Type*locality),*and*four*individuals*of*M.'t.' microgalbineus'(M200,*M209,*M448,*M449*[possibly*also*M431,*see*below]).*These* subspecies*are*differentiated*from*one*another*by*a*combination*of*scale*counts,* geography,*number*of*black*body*rings,*and*the*size*of*the*nuchal*ring*(Roze*1996;* Campbell*and*Lamar*2004;*LavinSMurcio*and*Dixon*2004).*Interestingly*three* subspecies,*M.*t.*fitzingeri,*M.*t.*maculatus,*and*M.*t.*microgalbineus,'occur*in*close* geographic*proximity*in*Mexico*and*are*thought*to*intergrade*(Fig.*1,*B).*We*sampled*all* Streicher et al. 3 four*subspecies*in*our*mitochondrial*analysis*and*none*of*them*was*found*to*be* monophyletic*(Fig.*1,*A).*We*sampled*M.'t.'maculatus,*M.'t.'microgalbineus,*and*M.'t.' tener*in*our*nuclear*phylogenetic*analysis*and*found*that*while*individuals*from*Mexico* possessed*more*private*alleles*(and*thus*longer*branch*lengths*in*general),*none*of*the* Mexican*subspecies*were*monophyletic*relative*to*the*other*subspecies*of*M.'tener'(Fig.* 1,*C).*However,*collectively*all*three*subspecies*of*M.'tener*were*found*to*be* monophyletic*relative*to*M.'fulvius.*Thus,*we*report*no*genetic*evidence*in*support*of* recognizing*the*subspecies*of*M.'tener*as*distinct*evolutionary*entities.** We*report*a*novel*locality*record*from*the*Mexican*state*of*Veracruz*for*a* specimen*that*possesses*mitochondrial*DNA*from*M.'tener*(M214,*Field*ID*JAC*22605;* Fig.*1;*A*and*B).*This*locality*was*included*in*Castoe*et*al.*(2012),*but*not*discussed*for* its*novelty*relative*to*previous*estimates*of*the*distribution*of*M.'bernadi*+*M.'tener'+* M.'tamaulipensis*(sensu*Campbell*and*Lamar*2004;*LavinSMurcio*and*Dixon*2004).*The* specimen*is*from*near*the*Los*Tuxtlas*mountain*range,*renowned*for*its*endemic* biodiversity*(PerezSHigareda*&*Navarro*1980).*We*initially*identified*specimen*M214*as* M.'diastema*given*the*collection*locality*(no*individuals*of*M.'tener'have*been*reported* near*the*Isthmus*of*Tehuantepec).*Interestingly,*Fraser*(1973)*noted*that*M.'diastema' from*Veracruz*had*fewer*ventral*and*subcaudal*scales*than*M.'diastema*populations* from*the*Yucatan*peninsula*(the*origin*of*our*outgroup*sample*of*M.'diastema,*M50;* Fig.*1,*A)*and*Guatemala/Honduras.*Thus,*although*we*lack*the*geographic*sampling*to* speculate*with*any*confidence,*our*discovery*of*an*individual*with*M.'tener* mitochondrial*DNA*from*an*area*where*M.'diastema*should*be*the*only*species*of* Streicher et al. 4 coralsnake*with*a*monadal*and*tricolored*dorsal*pattern*(M.'limbatus*and*M.'elegans' are*snakes*with*a*bicolor*and*triadal/pentadal*dorsal*pattern,*respectively),*warrants* future*investigation.* Based*on*our*mitochondrial*phylogenies*(Fig.*S2),*we*find*support*for*the*M.' fulvius*clade*(M.'tener*and*M.'fulvius)*being*sister*to*M.'nigrocinctus*(Girard,*1855),'a* species*with*notable*color*pattern*polymorphism*from*Central*America*and*Mexico*(see* Campbell*and*Lamar,*2004),*or*to*a*clade*of*M.'elegans'(Jan*1858)*and*M.'nigrocinctus* (Fig.*1,*A;*Fig*S2).*Micrurus'diastema,'M.'fulvius,*M.'nigrocinctus,*M.'tener,*belong*to*the* monadal*group*of*New*World*coralsnakes*(Slowinski*1995;*Campbell*and*Lamar*2004;* Renjifo*et*al.*2012).*Micrurus'elegans*and*M.'laticollaris*(Peters*1869)*belong*to*the* Central*American*triadSbearing*group.*Thus,*our*phylogenetic*analyses*of*mitochondrial* DNA*may*suggest*that*the*monadal*and*Central*American*triadal*color*groups*are*nonS monophyletic.* Justification(for(synonymizing(Micrurus'bernadi(with(M.'tener' Castoe*et*al.*(2012)*defined*the*Micrurus'fulvius*complex*as*including*M.'bernadi,*M.' fulvius,*M.'tener,*and*M.'tamaulipensis.*The*association*between*M.'tener*and*M.' fulvius'was*well*established*(see*Campbell*and*Lamar*2004)*and*the*close*phylogenetic* affinities*of*M.'tener*with*M.'tamaulipensis*were*suspected*(Lavin*Murcio*and*Dixon,* 2004;*next*section).*However,*the*inclusion*of*M.'bernadi*in*the*M.'fulvius*complex*was* largely*unprecedented*(Castoe*et*al.*[2012]*incorrectly*attributed*the*placement*to* LavinSMurcio*and*Dixon*[2004]).*Schmidt*(1933)*posited*that*it*was*“…possible*that*the* Streicher et al. 5 coloration*of*the*specimen,*in*which*the*black*rings*are*reduced*to*a*series*of*dorsal* spots,*is*an*individual*anomaly.”*Although*he*did*mention*that*the*holotype*of*M.' bernadi'was*the*northernmost*record*in*Mexico*for*a*coralsnake*outside*of*M.'tener,* Schmidt*(1933)*did*not*clearly*indicate*what*species*the*color*pattern*anomaly*would*be* referable*to.*In*contrast,*PérezSHigareda*and*Smith*(1990)*hypothesized*that*M.'bernadi* was*a*close*relative*of*M.'diastema,*which*also*occurs*in*eastScentral*Mexico.** Castoe*et*al.*(2012)*included*M.'bernadi*in*the*M.'fulvius*complex*because*a* single*individual*of*this*taxon*had*a*microsatellite*profile*that*clustered*with*M.'tener.* Although*Castoe*et*al.*(2012)*did*not*specifically*report*it,*this*individual*(M431,*Field*ID* ITAH*1189;*Table*S1;*Fig.*5)*was*identified*as*M.'bernadi*on*the*basis*of*the*possessing* the*distinctive*“saddled”*color*pattern*observed*in*the*holotype.*We*included*specimen* M431*in*our*study,*and*like*Castoe*et*al.*(2012)*found*with*microsatellites,*our*nuclear* and*mitochondrial*DNA*data*suggest*it*is*indeed*nested*within*M.'tener'(Fig.*1,*A*and*C).* Our*mitochondrial*analyses,*which*included*M.'diastema'as*an*outgroup,*further* supported*this*relationship*(Fig.*1,*A;*Fig.*S2).*However,*specimen*M431*was*collected* about*7*km*south*of*Huejutla*de*Reyes*in*Hidalgo,*which*is*over*50*km*north*of*the*type* locality*of*M.'bernadi*in*Zacualtipan,*Hidalgo.*The*collection*locality*of*specimen*M431* occurs*where*the*ranges*of*M.'t.'microgalbineus*and*M.'bernadi*putatively*overlap* (Campbell*and*Lamar*2004).*Distinguishing*these*taxa*morphologically*is*not* straightforward*as*scale*count*data*overlap*in*both*sexes*(condition*of*M.'t.' microgalbineus*[M.'bernadi*in*brackets]:*ventrals,*males*198S204*[198–212],*females* 216S225*[212–225];*subcaudals,*males*41–45*[45–48],*females*32–38*[34–39],*temporals* Streicher et al. 6 1+1*[1+1])*and*both*taxa*possess*color*pattern*polymorphism*(Schmidt*1958;*Roze* 1996;*Campbell*and*Lamar*2004).*Thus,*it*is*possible*that*specimen*M431*is*an* individual*of*M.'bernadi,*but*it*is*equally*likely*that*it*is*an*aberrant*color*morph*of*M.'t.' microgalbineus.*To*differentiate*between*these*possibilities,*we*sequenced* mitochondrial*DNA*from*three*additional*specimens*referable*to*M.'bernadi,*all* originating*from*near*Cuetzalan,*Puebla*(M201,*Field*ID*JAC*22468;*M236*Field*ID*ENS* 10590;*and*M246,*Field*ID*ENS*10790).*These*specimens*had*also*been*included*in*the* microsatellite*screening*portion*of*Castoe*et*al.*(2012;*their*Fig.*1,*A),*but*not*the* preliminary*population*genetic*analysis.*Cuetzalan*is*a*wellSknown*locality*for*M.' bernadi*that*is*included*in*the*species*account*provided*by*Campbell*and*Lamar*(2004).*
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