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).*
This*locality*is*only*40*km*straightSline*distance*from*Necaxa,*Puebla*another*locality* that*has*yielded*multiple*specimens*of*M.'bernadi*(Schmidt,*1958).*The*three*specimens* from*Cuetzalan*all*have*M.'tener*mitochondrial*haplotypes*(Fig.*1,*A),*although*they* belong*to*a*different*haplogroup*than*specimen*M431*from*Hidalgo.*Thus,*while*we* encourage*future*exploration*of*this*issue*via*genetic*comparisons*with*material*from* near*the*type*locality*of*M.'bernadi,'we*believe*our*genetic*data*and*a*lack*of* morphological*autapomorphies*warrant*synonymizing*M.'bernadi*with*M.'tener.***
Justification(for(synonymizing(Micrurus'tamaulipensis(with(M.'tener'
Micrurus'tamaulipensis*was*described*on*the*basis*of*four*specimens*collected*from* pine*oak*forest*in*a*small*region*of*Tamaulipas*known*as*the*Sierra*de*Tamaulipas.*In* the*original*description,*LavinSMurcio*and*Dixon*(2004)*indicated*phylogenetic*affinities* Streicher et al. 7 between*M.'tamaulipensis*and*M.'tener:*"On*comparison*to*the*only*other*species*of* coral*snake*in*the*region*(M.'tener),*it*is*obvious*that*the*latter*species*is*its*closest* relative."*Our*study*included*mitochondrial*DNA*from*a*specimen*collected*in*the*Sierra* de*Tamaulipas*(M326,*Field*ID*JAC*24615).*Importantly,*specimen*M326*is*from*near*
Acuña,*where*the*paratype*of*M.'tamaulipensis*was*collected*(UMMZ*95201).*We*found* that*mitochondrial*DNAs*placed*specimen*M326*as*nested*between*two*samples*of*M.' bernadi'from*Puebla*(Fig.*1,*A).*Morphological*similarities*to*the*two*subspecies*of*M.' tener*that*occur*closest*to*the*range*of*M.'tamaulipensis,*M.'t.'maculatus*and*M.'t.' microgalineus*(Fig.*1,*B),*were*noted*in*the*original*description*(LavinSMurcio*and*Dixon,*
2004).*Interestingly,*these*three*forms*occur*in*a*putative*zone*of*intergradation* between*subspecies*of*M.'tener*(Fig.*1,*B;*Campbell*and*Lamar,*2004).*Comparisons*of* morphology*among*snakes*from*this*region*suggest*overlap*in*characteristics*that*have* been*used*to*diagnose*species*and*subSspecies*(condition*of*M.'t.'maculatus,*M.'t.' microgalbineus,*and*M.'tamaulipensis,*respectively):*ventrals*in*males*185–195,*198–
204,*193–202;*ventrals*in*females*205–208,*216–225,*209;*subcaudals*in*males*43–45,*
41–45,*42–46;*subcaudals*in*females,*31,*32–38,*31;*temporals*1+1,*1+1,*1+1/2.*In*their* diagnosis*LavinSMurcio*and*Dixon*(2004)*state*that'M.'tamaulipensis'differs*from*M.' tener*by*the*presence*of*a*black*head*cap*that*extends*beyond*the*tip*of*the*parietal* scales*four*to*six*scales.*This*statement*is*misleading*because*they*are*actually*referring* to*the*width*of*the*first*black*band,*the*nuchal*band,*and*not*just*the*cap*as*they*state.*
Another*misleading*statement*in*the*diagnoses*is*that*M.'tamaulipenis'lacks*a*yellow* head*ring*across*the*interparietal*suture*(present*in*M.'tener),*which*could*be* Streicher et al. 8 interpreted*as*some*of*their*specimens*having*a*virtually*complete*yellow*head*ring,* except*for*the*suture.*The*extensive*black*cap*with*fusion*to*the*nuchal*ring*has*been* reported*for*individuals*of*M.'bernadi'(Roze*1996),*and*can*also*be*seen*in*the* population*of*M.'diastema*in*the*northeastern*Yucatan*Peninsula,*some*that*have* extensive*red*coloration,*sometimes*even*including*the*tail.*The*only*character*used*to* definitively*differentiate*M.'tamaulipensis*from*M.'tener*is*the*presence*of*three*colors* on*the*tail*(black,*yellow,*and*red).*Although*most*individuals*of*M.'tener*have*yellow* and*black*tails,*there*are*many*examples*of*specimens*with*red*suffusions*on*the*tails*
(typically*occurring*as*red*blotches*or*rings*in*the*center*of*yellow*rings;*E.N.*Smith*pers.* obs.).*Given*the*levels*of*color*pattern*polymorphism*known*to*occur*within*M.'tener,* the*utility*of*color*pattern*characters*for*differentiating*taxa*especially*in*Mexico*is* highly*subjective*and*possibly*dubious.*In*light*of*the*evidence*from*scale*counts*and* genetics*that*do*not*clearly*distinguish*it*from*M.'tener,*we*suggest*recognizing*M.' tamaulipensis*as*a*junior*synonym*of*M.'tener.*
Based*on*our*collective*findings,*we*present*below*a*revised*species*account*for*
M.'tener*including*new*synonymies,*diagnostic*comparisons*with*closely*related*and* sympatric*species,*an*updated*geographic*distribution,*and*comments*on*morphological* variation*in*this*widespread*species*of*venomous*snake.**
'
' Streicher et al. 9
Micrurus'tener((Baird(and(Girard,(1853)(
Figs.*1–6*and*S1–S9*
Elaps'tenere*Baird*and*Girard,*1853,*Cat.'N.'Am.'Rept.*1:172*[22,*156].*Syntypes:*Originals*lost,*
USNM*1121*designated*lectotype*(Roze,*1996).*Type*Locality:*New*Braunfels,*Texas.*
USA.*
Elaps'tristis*Baird*and*Girard,*1853,*Cat.'N.'Am.'Rept.*1:172*[23].*In*Part.*Syntypes:*USNM*1123*
(M'.tener)*and*1124*(M.'fulvius,'from*Mississippi).*Type*Locality:*Rio*Grande,*W*of*San*
Antonio,*Texas,*USA.*
Elaps'fitzingeri*Jan,*1858,*Rev.'Mag.'Zool.*10:514S527*[521,*Pl.*1].*Syntypes:*All*lost*except*for*
NMW*18297.*Type*Locality:*Guanajuato,*Mexico*(Smith*and*Taylor,*1950).*
Elaps'tener*–*Günther,*1859,*Proc.'Zool.'Soc.'London*1859:79S89*[86].*
Elaps'fulvius*var.*fitzingeri'–*Jan,*1863,*Elenco*Sist.*Ofidi*1S143*[113].*
Elaps'fulvius'tener*–*Cope,*1875,*Bull.'U.S.'Natl.'Mus.'1:1S104*[34].*
Elaps'bernadi*Cope,*in*FerrariSPerez,*1886,*Proc.'U.S.'Natl.'Mus.*9:125S199*[190].*[Nomen'
nudem]*syn.(nov.*
Elaps'bernadi*Cope,*1887,*Bull.'U.S.'Natl.'Mus.*32:1S98[87]*Holotype:*ANSP*14767,*Type*Locality:*
Zacualtipan,*Hidalgo,*Mexico.*syn.(nov.*
Elaps'fulvius*–*Boulenger,*1896,*Cat.'Snakes'British'Mus.*3:1S727*[422].*In*Part.*
Micrurus'bernadi*–Schmidt,*1933,*Field'Mus.'Nat.'Hist.'Publ.,'Zool.'Ser.*20:29S40*[40]*syn.(nov.*
Micrurus'fitzingeri*–*Schmidt,*1933,*Field'Mus.'Nat.'Hist.'Publ..'Zoo.'Ser.*20:29S40*[38].*
Micrurus'fulvius'tenere*–*Schmidt*1933,*Field'Mus.'Nat.'Hist.'Publ..'Zoo.'Ser.*20:29S40*[40].*
Micrurus'fitzingeri'fitzingeri*–*Brown*and*Smith,*1942,*Proc.'Biol.'Soc.'Washington*55:63S66*[63].*
Micrurus'fitzingeri'microgalbenius*Brown*and*Smith,*1942,*Proc.'Biol.'Soc.'Washington*55:63S66* Streicher et al. 10
[63].*Holotype:*Strecker*Museum*14984.*Type*Locality:*Antiguo*Morelos,*Tamaulipas,*
Mexico.*
Micrurus'fulvius'maculatus*Roze,*1967,*Am.'Mus.'Novitat.*2287:1S60*[27].'Holotype:*ZMH*5685.*
Type*Locality:*Tampico,*Tamaulipas,*Mexico.*
Micrurus'fulvius'microgalbineus*–*Roze,*1967,*Am.'Mus.'Novitat.*2287:1S60*[26].'
Micrurus'fulvius'fitzingeri*–*Roze,*1967,*Am.'Mus.'Novitat.*2287:1S60*[29].*
Micrurus'fulvius'tener**–*Frost*and*Collins,*1988,'Herpetol.'Rev.*19:73S74*[73].**
Micrurus'tener'–*Collins,*1991,*Herpetol.'Rev.*22:42S43*[43].*
Micrurus'diastema'bernadi*–*PérezSHigareda*and*Smith,*1990,*Bull.'Maryland'Herpetol.'Soc.'
26:5S13*[5].*syn(nov.*
Micrurus'tener'fitzingeri*–*Campbell*and*Lamar,*2004,*The'Venomous'Reptiles'of'the''Western'
Hemisphere'1:1S475*[195].*
Micrurus'tener*maculatus*–*Campbell*and*Lamar,*2004,*The'Venomous'Reptiles'of'the''Western'
Hemisphere'1:1S475*[196].**
Micrurus'tener*microgalbineus*–*Campbell*and*Lamar,*2004,*The'Venomous'Reptiles'of'the''
Western'Hemisphere'1:1S475*[195].*
Micrurus'tamaulipensis*LavinSMurcio*and*Dixon,*2004,*Phyllomedusa*3:3S7*[4].*Holotype:*ITT*
751.*Type*Locality:*Sierra*de*Tamaulipas,*Rancho*La*Sauceda,*Tamaulipas,*Mexico.*syn.(
nov.(
*
English(and(Spanish(names:(Texas*Coralsnake,*Coral*Tejano,*Coralillo*Tejano*
Distribution(and(habitat:*Across*a*variety*of*habitats*spanning*arid*to*high*elevation* regions*of*the*Central*Mexican*Plateau,*cloud*forests*of*the*Sierra*Madre*Oriental,*the*
Edwards*Plateau*region*of*central*Texas,*and*wooded*areas*of*Louisiana*and*east*Texas,* Streicher et al. 11
USA.*This*species*also*inhabits*tropical*wetSforests*of*the*Gulf*coast*of*Mexico*extending* as*far*south*as*the*Isthmus*of*Tehuantepec*(Fig.*1,*B).*Collection*localities*for*specimens* used*in*our*study*ranged*in*elevation*from*6*m*(Louisiana,*USA)*to*2155*m*(Guanajuato,*
Mexico)*above*sea*level.**
Description:'Adult*body*sizes*that*can*range*from*550–1181*mm*but*is*generally*less* than*800*mm*total*length.*Ventral*scale*counts*range*from*185–216*in*males*and*205–
231*in*females.*Subcaudal*scale*counts*range*from*38–48*in*males*and*26–41*in*females.*
Temporal*scales*can*have*either*the*condition*1+1*or*1+2.*There*are*between*10*and*27* black*body*rings*(although*not*always*complete)*in*males,*and*10*to*26*in*females.*
There*are*between*3–12*tail*rings*in*males*and*2–8*tail*rings*in*females.**
Similar(species:(The*closest*extant*relative*of*M.'tener*is*likely*M.'fulvius*(Fig.*1,*A*and*
C).*These*taxa*can*be*differentiated*by*the*condition*of*the*black*nuchal*band*where*it* does*not*reach*the*tips*of*the*parietal*scales*in*M.'fulvius*(where*it*does*in*M.'tener,'but* is*obscured*by*a*black*nuchal*cap*in*specimens*from*the*Sierra*de*Tamaulipas).*The* extent*that*this*band*extends*onto*the*parietals*varies*across*populations*of*M.'tener*
(as*does*the*relative*completeness*of*the*parietal*light*ring;*e.g.*Campbell*and*Lamar,*
2004,*their*Fig.*26;*LavinSMurcio*and*Dixon,*2004,*their*Fig.*1).*In*Mexico,*the*only*other* coralsnakes*that*possibly*have*overlapping*ranges*with*M.'tener*are*M.'browni,' diastema,'M.'limbatus*and*M.'elegans.*Micrurus'elegans*has*black*body*rings*disposed* in*triads*or*pentads*alternating*over*white*rings*and*between*orange*rings*(M.'tener*is* monadal*or*bicolored*throught*the*body).*Micrurus'limbatus*is*a*bicolored*red*
(orange)/black*snake,*that*either*has*black*saddles*or*narrow*rings*over*a*red*or*dark* Streicher et al. 12 orange*body.*Both*M.'browni*and*M.'diastema*also*have*body*dorsa*with*a*highly* variable*number*of*black*rings,*10–27*and*0–62,*respectively.*M.'diastema*has* populations*with*the*body*dorsum*bicolored*or*tricolored*and*monadal,*and*both* species*can*have*very*narrow*yellow*rings.*Both*species*overlap*and*surpass*in*meristic* characters*the*variation*seen*in*M.'tener,*although*these*two*species*also*need*to*be* reevaluated*taxonomically*based*on*our*extension*of*the*M.'tener*distribution*into*
Veracruz,*and*they*both*contain*several*distinctly*recognizable*subspecies*in*need*of* molecular*evaluation.*In*general*M.'tener*in*Mexico*has*a*black*mental*scale*and*black* anterior*infralabials*(first*three),*dark*pigmentation*over*the*chinSshields,*and*black* color*extending*widely*into*the*gular*area.*In*M.'diastema*from*north*of*the*Isthmus*of*
Tehuantepec*the*black*color*on*the*infralabials*and*mental*scale*tends*to*be*reduced*or* absent,*the*chin*tends*to*be*relatively*free*of*dark*color,*and*the*gular*area*tends*to*be* relatively*reduced*of*black*coloration,*at*least*medially*and*just*behind*the*head.*
Micrurus'browni'is*very*similar*in*color*to*M.'tener,*but*adult*males*and*even*some*large* adult*females*of*this*species*differ*from*both*M.'tener*and*M.'diastema*in*having* supracloacal*tubercles.*
Color(pattern(polymorphism:*Complex*variation*in*color*pattern*has*been*widely* documented*in*the*monadal*group*of*Micrurus*(Savage*and*Slowinski*1992).*Varying* degrees*of*melanistic*coloration*are*known*to*occur*in*M.'tener*from*Texas;*sometimes* resulting*in*envenomation*based*on*misidentification*(Gloyd*1938;*Campbell*and*Lamar*
2004).*This*melanistic*phenotype*is*particularly*abundant*around*San*Antonio,*Texas,*
USA*(Werler*and*Darling*1950).*Interestingly,*our*revised*concept*of*M.'tener*increases* Streicher et al. 13 the*levels*of*color*pattern*polymorphism*known*from*the*species*(to*include*black* heads*and*nonSringed*saddle*patterns).*The*types*of*color*pattern*polymorphism* observed*in*M.'tener*are*strikingly*similar*to*those*observed*in*several*species*of* colubrid*mimics*inhabiting*similar*regions*of*Mexico*(e.g.*Sonora'aemula*(Cope*1879),*S.' michoacanensis'(Dugès*1884),*S.'mutabilis'Stickel*1943,*Cox*et*al.*2012;*Pliocercus' elapoides'Cope*1860,'Campbell*and*Lamar*2004).*Interestingly,*these*mimics*include* multiple*(presumably*independent)*instances*of*evolving*the*“saddle”*that*also*occurs*in* populations*of*M.'tener'previously*referred*to*as*M.'bernadi.**
Conclusions(and(future(directions(
In*his*review*of*the*genus*Micrurus,*Karl*Schmidt*(1933)*wrote,*“The*genus*Micrurus*is* characterized*by*great*uniformity*of*arrangement*of*the*head*shields*and*the*number*of* dorsal*scale*rows.*It*becomes,*therefore,*the*more*necessary*to*scrutinize*the* taxonomic*characters*available.*One*of*the*first*results*of*this*critique*is*to*affirm*the* constancy*and*value*of*the*color*pattern*characters.”*Despite*this*recommendation,* much*of*the*subsequent*taxonomy*of*Micrurus*has*been*based*on*using*presumed* discrete*color*pattern*characters.*Given*the*findings*of*our*study,*we*strongly* recommend*that*data*sources*beyond*color*pattern*and*scale*counts*(e.g.*DNA,*viscera,* osteology,*and*dentition)*are*utilized*in*future*endeavors*to*untangle*evolutionary* relationships*among*Mexican*snakes*of*the*genus*Micrurus.*
* Streicher et al. 14
LITERATURE(CITED(
*
Boulenger,*G.*A.*1896.*Catalogue*of*the*snakes*at*the*British*Museum*(Natural*History).*
Vol.*3:727*pp.*
Campbell,*J.*A.,*and*W.*W.*Lamar.*2004.*The*Venomous*Reptiles*of*the*Western*
Hemisphere.*Cornell*University*Press,*Ithaca,*NY.*Vol.*1:475*pp.**
*
CarbajalSSaucedo,*A.,*E.*LópezSVera,*M.*BénardSValle,*E.*N.*Smith,*F.*Zamudio,*A.*R.*de*
Roodt*and*A.*OlveraSRodríguez.*2013.*Isolation,*characterization,*cloning*and*
expression*of*an*alphaSneurotoxin*from*the*venom*of*the*Mexican*coral*snake*
Micrurus*laticollaris*(Squamata:*Elaidae).*Toxicon*66:64–74.**
*
Castoe,*T.*A.,*E.*N.*Smith,*R.*M.*Brown*and*C.*L.*Parkinson.*2007.*HigherSlevel*phylogeny*
of*Asian*and*American*coralsnakes,*their*placement*within*the*Elapidae*(Squamata),*
and*the*systematic*affinities*of*the*enigmatic*coralsnake*Hemibungarus'calligaster.*
Zool.*J.*Linn.*Soc.*151:809–831.***
Castoe,*T.*A.,*J.*W.*Streicher,*J.*M.*Meik,*M.*J.*Ingrasci,*A.*W.*Poole,*A.*P.*J.*de*Koning,*J.*
A.*Campbell,*C.*L.*Parkinson,*E.*N.*Smith,*and*D.*D.*Pollock.*2012.*Thousands*of*
microsatellite*loci*from*the*venomous*coralsnake*Micrurus'fulvius*and*variability*of*
select*loci*across*populations*and*related*species.*Mol.*Ecol.*Resour.*12:1105–1113* Streicher et al. 15
*
Cox,*C.*L.,*A.*R.*Davis,*J.*ReyesSVelasco,*P.*PonceSCampos,*E.*N.*Smith,*O.*FloresSVIllela*
and*J.*A.*Campbell.*2012.*Molecular*systematics*of*the*genus*Sonora*(Squamata:*
Colubridae)*in*central*and*western*Mexico.*Syst.*Biodivers.*10:93–108.**
*
Fraser,*D.*F.*1973.*Variation*in*the*coral*snake*Micrurus'diastema.*Copeia*1973:1–17.*
*
Gloyd,*H.*K.*1938.*A*case*of*poisoning*from*the*bite*of*a*black*coral*snake.*Herpetologica*
1:121–124.*
LavinSMurcio,*P.*A.*and*J.*R.*Dixon.*2004.*A*new*species*of*coral*snake*(Serpentes,*
Elapidae)*from*the*Sierra*de*Tamaulipas,*Mexico.*Phyllomedusa*3:3–8.*
*
Margres,*M.J.,*K.*Aronow,*J.*Loyacano*and*D.*R.*Rokyta.*2013.*The*venomSgland*
transcriptome*of*the*eastern*coral*snake*(Micrurus'fulvius)*reveals*high*venom*
complexity*in*the*intragenomic*evolution*of*venoms.*BMC*Genomics*14:531.**
*
PerezSHigareda,*G.*and*L.*D.*Navarro.*1980.*The*faunistic*districts*of*the*low*plains*of**
******Veracruz,*Mexico,*based*on*reptilian*and*mammalian*data.*Bull.*Md.*****
******Herpetol.*Soc.*16:54–69.**
*
PerezSHigareda,*G.*and*H.*M.*Smith.*1990.*The*endemic*coral*snakes*of*the*Los*Tuxtlas** Streicher et al. 16
*******region,*southern*Veracruz,*Mexico.*Bulletin*of*the*Maryland*Herpetological*Society***
*******26:5–13.***
*
Pyron,*R.A.,*F.*T.*Burbrink,*G.*R.*Colli,*A.*NietoSMontes*de*Oca,*L.*J.*Vitt,*C.*A.*Kuczynski**
*****and*J.*J.*Wiens*2011.*The*phylogeny*of*advanced*snakes*(Colubroidea),*with*the**
****discovery*of*a*new*subfamily*and*comparison*of*support*methods*for*likelihood*trees.**
*****Mol.*Phylogenet.*Evol.*58:329–342.**
*
Pyron,*R.A.,*F.*T.*Burbrink*and*J.*J.*Wiens.*2013.*A*phylogeny*and*revised*classification*of**
******Squamata,*including*4161*species*of*lizards*and*snakes.*BMC*Evol.*Biol.**
******13:93.**
*
Renjifo,*C.,*E.*N.*Smith,*W.*C.*Hodgson,*J.*M.*Renjifo,*A.*Sanchez,*R.*Acosta,*J.*H.**
*****Maldonado*and*A.*Riveros.*2012.*Neuromuscular*activity*of*the*venoms*of*Columbian**
****coral*snakes*Micrurus'dissoleucus*and*Micrurus'mipartitus:*An*evolutionary**
*****perspective.*Toxicon*59:132–142.**
*
Roze,*J.*A.*1996.*Coral*snakes*of*the*Americas:*Biology,*identification,*and*venoms.*
Malabar,*Florida.*Krieger*Publishing,*340*pp.**
*
Savage,*J.*M.*and*J.*B.*Slowinski.*1992.*The*colouration*of*the*venomous*coral*snakes**
******(family*Elapidae)*and*their*mimics*(families*Aniliidae*and*Colubridae).*Biol.* Streicher et al. 17
******J.*Linn.*Soc.*45:*235–254.**
*
Schmidt,*K.*P.*1933.*Preliminary*account*of*the*coral*snakes*of*Central*America*and*
Mexico.*Field*Mus.*Nat.*Hist.*Zool.*Ser.*20:29–40.*
*
Schmidt,*K.P.*1958.*Some*rare*or*littleSknown*Mexican*coral*snakes.*Fieldiana*Zool.*
39:201–212.**
*
Slowinski,*J.B.*1995.*A*phylogenteic*analsis*of*the*New*World*coral*snakes*(Elapidae:*
Leptomicrurus,*Micruriodes,*and*Micrurus)*based*on*allozymic*and*morphological*
characters.*J.*Herpetol.*29,*325–338.**
*
Tanaka,*G.D.,*M.*de*Fátima*D.*Furtado,*F.*C.*V.*Portaro,*O.*A.*Sant'Anna*and*D.*V.*
Tambourgi.*2010.*Diversity*of*Micrurus**snake*species*related*to*their*venom*toxic*
effects*and*the*prospective*of*antivenom*neutrilization.*PLoS*Negl.*Trop.*Dis.*4:e622.**
*
Werler,*J.*E.*and*D.*M.*Darling.*1950.*A*case*of*poisoning*from*the*bite*of*a*coral*snake,*
Micrurus'f.'tenere*Baird*and*Girard.*Herpetologica*6:197–199.*
*
Appendix(S2:(Additional(Methods(
RADSEQ(LIBRARY(CONSTRUCTION(AND(SEQUENCING(
DNA$isolates$were$quantified$using$a$Qubit$fluorometer$and$dsDNA$HS$assay$kit$(Life$
Technologies),$and$approximately$250$ng$of$DNA$for$each$individual$with$the$ restriction$enzymes$SbfI$and$Sau3AI)(New$England$Biolabs).$Following$adaptor$ ligation,$magneticNbead$purified$samples$were$pooled$and$size$selected$for$ fragments$ranging$from$440$and$540$bp$using$a$1.5%$agarose$gel$cartridge$run$on$a$
Blue$Pippin$Prep$(Sage$Science).$RAD$libraries$were$amplified$via$PCR$with$Phusion®$
DNA$polymerase$(New$England$Biolabs),$purified$using$magnetic$beads,$and$ quantitated$using$a$DNA$7500$chip$on$a$Bioanalyzer$2100$(Agilent$Technologies).$
Amplified$libraries$were$pooled$and$sequenced$on$a$single$Illumina$HiSeq$2500$lane$ using$100$bp$pairedNend$reads.$
ESTIMATING(THE(HISTORICAL(RANGE(OF(CORALSNAKES(
We$modeled$the$climatic$niche$of$M.)fulvius)and)M.)tener$to$approximate$the$ current$and$last$glacial$maximum$(LGM)$distribution$of$each$species.$We$applied$an$ ecological$niche$modeling$method,$where$environmental$data$are$extracted$from$ current$occurrence$records,$and$habitat$suitability$is$evaluated$across$the$landscape$ using$a$programNspecific$algorithm$(Elith$et$al.$2006).$The$presentNday$models$of$ each$species$were$then$projected$on$the$climatic$reconstructions$of$the$LGM$under$ the$assumption$that$the$climatic$niche$of$each$species$remained$conserved$between$ the$LGM$and$present$(Elith$et$al.$2010).$$
For$occurrence$data,$we$used$museum$records$downloaded$from$the$Vertnet$ Database$(http://vertnet.org/)$and$our$own$observations.$Our$datasets$comprised$
176$records$for$M.)fulvius$and$128$records$for$M.)tener.$The$current$climate$was$ represented$by$bioclimatic$variables$from$the$WorldClim$dataset$v.$1.4$(Hijmans$et$ al.$2005)$with$resolution$of$2.5$minutes.$We$followed$the$methodology$of$Jezkova$et) al.$(2011)$to$remove$highly$correlated$variables$(i.e.$with$a$correlation$coefficient$>$
0.9),$resulting$in$selection$of$12$predictor$variables$(out$of$the$19$bioclimatic$ variables).$For$environmental$layers$representing$the$climatic$conditions$of$the$LGM,$ we$used$three$models$of$oceanNatmosphere$simulations:$CCSM4,$MIROCNESM,$and$
MPINESMNP$(Braconnot$et$al.$2007),$all$available$through$www.wordclim.org.$$
Climatic$niche$models$were$constructed$in$MAXENT$v.$3.3.3k$(Phillips$et$al.$2008),$ which$estimates$relative$probabilities$of$the$presence$of$species$within$defined$ geographic$spaces,$with$high$probabilities$indicating$suitable$environmental$ conditions$for$the$species$(Phillips$et$al.$2004).$Our$models$were$confined$to$the$ southeastern$region$of$North$America.$We$used$the$default$parameters$in$MAXENT$
(500$maximum$iterations,$convergence$threshold$of$0.00001,$regularization$ multiplier$of$1,$and$10,000$background$points)$with$crossNvalidation$used$as$a$ replicated$run$type.$We$removed$duplicate$presence$records$(resulting$in$a$final$ dataset$of$139$records$for$M.)fulvius$and$115$records$for$M.)tener).$We$ran$20$ replicates$for$each$model,$and$an$average$model$was$presented$using$logistic$ probability$classes$of$climatic$niche$suitability.$The$presenceNabsence$maps$were$ determined$using$a$threshold$that$balances$training$omission,$predicted$area$and$ threshold$value$(less$conservative)$and$a$threshold$that$equates$entropy$of$ threshold$and$original$distributions$(more$conservative).$We$used$the$receiver$ operating$characteristic$to$determine$an$area$under$the$curve$(AUC)$value$to$
evaluate$model$performance,$where$AUC$values$range$from$0.5$for$a$random$
prediction$to$1$for$perfect$prediction$(Raes$and$ter$Steege$2007).$
We$found$the$following$AUC$values$for$our$SDMs:$M.)fulvius,$0.967;$M.)tener,$0.928;$
M.$fulvius$complex$–$0.923.$The$less$and$more$conservative$thresholds$corresponded$
to$0.05$and$0.16$logistic$probabilities,$respectively.(
( LITERATURE(CITED(
$
Braconnot,$P.$et$al.$2007.$Results$of$PMIP2$coupled$simulations$of$the$MidNHolocene$
and$Last$Glacial$Maximum$N$Part$1:$experiments$and$largeNscale$features.$Clim.$
Past$3:261–277.$
$
Elith,$J.$et$al.$2006.$Novel$methods$improve$prediction$of$species'$distributions$from$
occurrence$data.$Ecography$29:129–151.$
$
Elith,$J.$et$al.$2010.$The$art$of$modelling$rangeNshifting$species.$Methods$Ecol.$Evol.$
1:330–342.$
$
Hijmans,$R.$J.$et$al.$2005.$Very$high$resolution$interpolated$climate$surfaces$for$
global$land$areas.$Int.$J.$Climatol.$25:1965–1978.$
$
Jezkova,$T.,$V.$OlahNHemmings$and$B.$R.$Riddle.$2011.$Niche$shifting$in$response$to$
warming$climate$after$the$last$glacial$maximum:$inference$from$genetic$data$and$ niche$assessments$in$the$chiselNtoothed$kangaroo$rat$(Dipodomys)microps).$Glob.$
Change$Biol.$17:3486–3502.$
$
Phillips,$S.$J.$and$M.$Dudik.$2008.$Modeling$of$species$distributions$with$Maxent:$new$
extensions$and$a$comprehensive$evaluation.$Ecography$31:161–175.$
$
Phillips,$S.$J.$et$al.$2004.$A$maximum$entropy$approach$to$species$distribution$
modeling.$Proceedings$of$the$twentyNfirst$international$conference$on$Machine$
learning.$
$
Raes,$N.$and$ter$H.$Steege.$2007.$A$nullNmodel$for$significance$testing$of$presenceN
only$species$distribution$models.$Ecography$30:727–736.$
$
$
Streicher(et(al.(
Table S1 Taxonomic and geographic sampling of the Micrurus fulvius complex. *For additional information related to voucher specimens and specimen locations, contact [email protected], [email protected], or [email protected].
No. Lab Taxon Locality (State: Country) GPS Latitude GPS Longitude GenBank* SRA
ID* (cyt-b/ND4) ddRADseq
1 M431 bernadi( Hidalgo: Mexico 21.083957 -98.458886 KU754310/ SAMN
KU754422 03451700
2 M236 bernadi( Puebla: Mexico 20.020327 -97.457428 KU754341/ None
KU754401
3 M246 bernadi( Puebla: Mexico 20.020327 -97.457428 KU754293/ None
None
4 M201 bernadi( Puebla: Mexico 20.063 -97.475 KU754350/ None
KU754400
5 M86 fulvius Tampa: Florida: USA 28.08971 -82.320659 KU754348/ SAMN
KU754436 03451693
6 M87 fulvius Walton: Florida: USA 30.687148 -86.110727 KU754345/ SAMN
KU754420 03451692
7 M89 fulvius Perry: Mississippi: USA 31.202179 -89.03775 KU754353/ None
KU754454 Streicher(et(al.(
8 M91 fulvius Lake: Florida: USA 28.723104 -81.474474 KU754333/ None
KU754439
9 M92 fulvius Highlands: Florida: USA 27.358501 -81.438075 KU754356/ None
KU754404
10 M172 fulvius Jefferson: Florida: USA 30.418585 -83.786444 KU754358/ None
None
11 M173 fulvius Highlands: Florida: USA 27.602164 -81.505093 KU754346/ SAMN
KU754440 03451688
12 M174 fulvius Broward: Florida: USA 26.286831 -80.24383 KU754309/ None
KU754430
13 M175 fulvius Liberty: Florida: USA 30.277667 -84.99125 KU754355/ None
KU754444
14 M176 fulvius Aiken: South Carolina: USA 33.290019 -81.703565 KU754303/ None
KU754410
15 M278 fulvius Palm Beach: Florida: USA 26.381072 -80.088876 KU754305/ SAMN
KU754395 03451689
16 M314 fulvius Franklin: Florida: USA 29.970327 -84.499021 KU754307/ SAMN
KU754425 03451691
17 M668 fulvius Orange: Florida: USA 28.599702 -81.196276 KU754365/ None
None Streicher(et(al.(
18 M691 fulvius Robeson: North Carolina: USA 34.625084 -79.016505 KU754363/ None
None
19 M736 fulvius Florida: USA (No other data) N/A N/A KU754373/ SAMN
KU754384 03451690
20 M692 fulvius New Hanover: North Carolina: USA 34.047341 -77.908151 KU754364/ SAMN
None 03451694
21 M51 tener fitzingeri( Querétaro: Mexico (No other data) N/A N/A KU754326/ None
KU754417
22 M516 tener fitzingeri( Guanajuato: Mexico 21.006016 -100.794708 KU754296/ None
None
23 M432 tener fitzingeri( Guanajuato: Mexico (No other data) N/A N/A None/ None
KU754427
24 M206 tener maculatus( Tamaulipas: Mexico 22.941543 -97.994608 KU754350/ SAMN
KU754400 03451696
25 M332 tener maculatus( Tamaulipas: Mexico (No other data) N/A N/A KU754330/ N/A
KU754396
26 M448 tener Hidalgo: Mexico 21.061049 -98.499485 KU754295/ SAMN microgalbineus( KU754435 03451699
27 M449 tener Hidalgo: Mexico 21.013163 -98.841985 KU754294/ SAMN microgalbineus( KU754431 03451698 Streicher(et(al.(
28 M200 tener Hidalgo: Mexico 21.003333 -98.336389 KU754314/ SAMN microgalbineus( KU754411 03451697
29 M209 tener Hidalgo: Mexico 21.163594 -98.400016 KU754360/ None microgalbineus( None
30 M88 tener tener Bienville: Louisiana: USA 32.34127 -93.107748 KU754332/ SAMN
KU754428 03451722
31 M90 tener tener Acadia: Louisiana: USA -92.39855 KU754331/ None
KU754426
32 M93 tener tener( Texas: USA (No other data) N/A N/A KU754334/ None
KU754406
33 M177 tener tener Val Verde: Texas: USA 29.413651 -100.904984 KU754352/ None
KU754432
34 M203 tener tener( Nuevo Leon: Mexico 25.573385 -100.225263 KU754354/ None
KU754418
35 M204 tener tener( Nuevo Leon: Mexico 25.613917 -100.357352 KU754342/ None
KU754424
36 M207 tener tener( Nuevo Leon: Mexico 25.589064 -100.200149 KU754359/ None
None
37 M208 tener tener( Nuevo Leon: Mexico 25.375004 -100.116565 KU754338/ None
KU754429 Streicher(et(al.(
38 M230 tener tener( Anderson: Texas: USA 31.936164 -95.888154 KU754337/ SAMN
KU754441 03451721
39 M267 tener tener( McMullen: Texas: USA 28.171798 -98.68361 KU754349/ None
KU754407
40 M268 tener tener( Travis: Texas: USA 30.308801 -97.591951 KU754343/ None
KU754452
41 M269 tener tener( McMullen: Texas: USA 28.502355 -98.540417 KU754328/ SAMN
KU754403 03451704
42 M270 tener tener( Travis: Texas: USA 30.397462 -97.682358 KU754371/ SAMN
KU754382 03451709
43 M275 tener tener( Bandera: Texas: USA 29.796669 -99.575714 KU754300/ None
KU754443
44 M279 tener tener( Brazos: Texas: USA 30.536885 -96.195376 KU754324/ SAMN
KU754405 03451720
45 M281 tener tener( La Salle: Texas: USA 28.254351 -99.046014 KU754316/ None
KU754412
46 M282 tener tener( Bandera: Texas: USA 29.806665 -99.568313 KU754327/ None
KU754448
47 M283 tener tener( Kendall: Texas: USA 29.9835 -98.603767 KU754340/ SAMN
KU754455 03451719 Streicher(et(al.(
48 M313 tener tener( Cameron: Texas: USA 25.854389 -97.396917 KU754357/ None
KU754442
49 M315 tener tener( Hays: Texas: USA 29.865052 -97.913451 KU754302/ None
KU754415
50 M316 tener tener( Travis: Texas: USA 30.302958 -97.936654 KU754318/ SAMN
KU754433 03451708
51 M320 tener tener( Milam: Texas: USA 30.607563 -97.218638 KU754306/ None
KU754398
52 M322 tener tener( Milam: Texas: USA 30.607563 -97.218638 KU754329/ None
KU754453
53 M323 tener tener( Travis: Texas: USA 30.415551 -97.790336 KU754320/ None
KU754413
54 M324 tener tener( Travis: Texas: USA 30.388172 -97.672553 KU754301/ None
KU764451
55 M325 tener tener( Jim Hogg: Texas: USA 27.247549 -98.823136 KU754319/ SAMN
KU754450 03451701
56 M334 tener tener( Dallas: Texas: USA 32.614075 -96.984918 KU754312/ None
KU754447
57 M342 tener tener( Jim Hogg: Texas: USA 27.277727 -98.662867 KU754321/ SAMN
KU754409 03451702 Streicher(et(al.(
58 M343 tener tener( Montgomery: Texas: USA 30.495532 -95.639323 KU754361/ None
None
59 M344 tener tener( Harris: Texas: USA 30.073333 -95.585833 KU754308/ SAMN 03451733 KU754445
60 M347 tener tener( Polk: Texas: USA 30.695493 -94.728014 KU754311/ None
KU754416
61 M348 tener tener( Travis: Texas: USA 30.179401 -97.725296 KU754315/ SAMN
KU754438 03451715
62 M426 tener tener( Montgomery: Texas: USA 28.332778 -98.066111 KU754323/ None
KU754437
63 M428 tener tener( Montgomery: Texas: USA 30.071944 -95.582222 KU754322/ None
KU754399
64 M445 tener tener( Jim Wells: Texas: USA 27.784172 -98.046518 KU754299/ SAMN
KU754456 03451703
65 M447 tener tener( Nueces: Texas: USA 27.683755 -97.743102 None/ SAMN
KU754419 03451705
66 M453 tener tener( Coahuila: Mexico 26.914882 -102.013811 KU754292/ SAMN
None 03451695
67 M574 tener tener( Walker: Texas: USA 30.16574 -96.06663 KU754298/ SAMN
None 03451732 Streicher(et(al.(
68 M575 tener tener( Starr: Texas: USA 30.235296 -96.365157 KU754336/ None
KU754397
69 M577 tener tener( Washington: Texas: USA 30.109131 -97.282754 KU754368/ None
None
70 M578 tener tener( Harris: Texas: USA 26.720055 -98.521407 KU754335/ None
KU754423
71 M583 tener tener( Menard: Texas: USA 30.913764 -99.857369 KU754362/ SAMN
None 03451714
72 M674 tener tener( Jasper: Texas: USA 31.047474 -94.308195 KU754370/ SAMN
None 03451731
73 M675 tener tener( Angelina: Texas: USA 31.333668 -94.191575 KU754369/ None
None
74 M676 tener tener( Jasper: Texas: USA 31.047474 -94.308195 KU754366/ SAMN
None 03451730
75 M703 tener tener( Goliad: Texas: USA 28.64176 -97.35498 KU754367/ SAMN
None 03451706
76 M735 tener tener Travis: Texas: USA (No other data) N/A N/A KU754372/ SAMN
KU754383 03451717
77 M738 tener tener( Harris: Texas: USA 30.075534 -95.583396 KU754374/ SAMN
KU754385 03451729 Streicher(et(al.(
78 M740 tener tener( Harris: Texas: USA 30.075534 -95.583396 KU754375/ SAMN
KU754386 03451718
79 M741 tener tener( Harris: Texas: USA 30.001392 -95.290115 KU754376/ SAMN
None 03451728
80 M742 tener tener( Harris: Texas: USA 30.001392 -95.290115 KU754377/ SAMN
KU754387 03451727
81 M743 tener tener( Harris: Texas: USA 30.556293 -95.789388 KU754378/ SAMN
KU754388 03451726
82 M744 tener tener( Montgomery: Texas: USA 30.550902 -95.80321 None/ SAMN
KU754389 03451713
83 M746 tener tener( Montgomery: Texas: USA 30.192755 -95.590108 None/ SAMN
KU754390 03451712
84 M747 tener tener( Montgomery: Texas: USA 30.075534 -95.583396 KU754379/ SAMN
KU754391 03451716
85 M748 tener tener( Harris: Texas: USA 30.55258 -95.750158 KU754380/ SAMN
KU754392 03451725
86 M749 tener tener( Montgomery: Texas: USA 29.999002 -95.20071 KU754290/ SAMN
None 03451724
87 M753 tener tener( Montgomery: Texas: USA 30.471216 -95.752187 KU754381/ SAMN
KU754393 03451723 Streicher(et(al.(
88 M754 tener tener( Montgomery: Texas: USA 30.471216 -95.752187 None/ SAMN
None 03451711
89 M755 tener tener( Brazos: Texas: USA 30.41942 -96.107476 KU754291/ SAMN
KU754394 03451707
90 M326 tamaulipensis( Tamaulipas: Mexico 23.21539 -98.45056 KU754304/ None
KU754402
91 M214 “diastema”( Veracruz: Mexico 18.331527 -95.092779 KU754297/ None
None
92 M33 laticollaris Morelos: Mexico N/A N/A KU754347/ None
KU754446
93 M15 browni Tabasco: Mexico N/A N/A KU754313/ None
KU754414
94 M50 diastema Quintana Roo: Mexico N/A N/A KU754325/ None
KU754434
95 M22 elegans Huehuetenango: Guatemala N/A N/A KU754339/ None
KU754449
96 M13 nigrocinctus Zacapa: Guatemala N/A N/A KU754344/ None
KU754421
97 M28 nigrocinctus Santa Rosa: Guatemala N/A N/A KU754317/ None
KU754457 Streicher(et(al.(
Table S2. Best nucleotide model scheme identified by Partition Finder. This scheme was used to conduct Bayesian phylogenetic analyses of concatenated mitochondrial DNA sequences (cytochrome-b and ND4).
Locus Sites Model Likelihood BIC
Cyt b pos 1 237 HKY+G -4547.85 10396.55
237 -4547.85 Cyt b pos 2 HKY +I 10396.55
Cyt b pos 3 236 GTR+G -4547.85 10396.55
ND4 pos 1 222 HKY+G -4547.85 10396.55
ND4 pos 2 222 HKY+I -4547.85 10396.55
ND4 pos 3 221 GTR+G -4547.85 10396.55
Streicher(et(al.(
Table S3. Number of RAD tags (unique restriction digestion loci) obtained from Illumina PE100 sequencing of the Micrurus fulvius complex allowing for different levels of percent missing individuals per SNP.
Dataset Coverage (Stack depth) Missing data No. RAD tags
Read 1 10X 50% 12,914
Read 1 20X 25% 1356
Read 2 10X 50% 7020
Read 2 20X 25% 450
Streicher(et(al.(
Table S4. Number of nuclear genome-wide SNPs obtained from the Micrurus fulvius complex. Bold indicates the SNP dataset that were used in the main text.
Dataset Coverage Missing No. SNPs Read No. SNPs Read Concatenated No. of 1 2 SNPs
Fixed SNPs 10X 50% 2138 1328 3466
Fixed SNPs 20X 25% 160 41 201
Biallelic 10X 50% 13,858 8689 22,547
Biallelic 20X 25% 1328 426 1754
Streicher(et(al.(
Table S5 Correlation coefficients (Spearman’s rho) for comparisons between latitude, longitude, and spatial principal components from Micrurus tener.
Correlation coefficient (rho) Correlation coefficient (rho)
Micrurus tener (all, N = 36) Micrurus tener (USA only, N = 30)
Spatial principal Latitude Longitude Latitude Longitude component
sPC 1 -0.48 -0.50 -0.21 -0.40 sPC 2 0.18 -0.11 0.38 0.60 sPC 3 0.49 0.72 -0.63 0.08 sPC 4 0.49 0.21 -0.18 -0.02 sPC 5 0.30 0.05 -0.07 0.11 sPC 6 0.00 0.18 0.09 -0.02 Streicher(et(al.( sPC 7 0.13 -0.05 -0.05 0.07 sPC 8 0.03 0.17 -0.03 0.17 sPC 9 0.05 0.16 -0.04 -0.04 sPC 10 0.09 0.08 -0.03 0.08
( Streicher et al. 1
Supporting*Figure*Legends*
Figure*S1*(Page*39).!Relationships!between!percent!missing!data!and!principal!component! scores!from!the!most!explanatory!axis!of!the!multivariate!nuclear!DNA!SNP!analysis.!
Figure*S2*(Page*40).!Bayesian!phylograms!of!cytochrome@b!(cyt@b)!and!NADH!dehydrogenase! subunit!4!(ND4)!generated!using!MrBayes.!Black!circles!on!nodes!correspond!to!posterior! probabilities!>!0.90.!
Figure*S3*(Page*41).!Results!of!k@means!clustering!in!adegenet!(Jombart!2008)!using!a! maximum!of!40!clusters!(30!retained!for!figure)!on!the!22,547!biallelic!nuclear!SNP!dataset!from!
45!individuals!of!the!Micrurus(fulvius!complex.!Note!inflection!point!between!two!and!four! clusters!indicated!by!arrow!indicating!optimal!number!of!clusters.!!
Figure*S4*(Page*42).!STRUCTURE!analysis!(K=3)!of!Micrurus(tener.!The!expected! heterozygosities!of!each!cluster!are!displayed!as!they!correspond!to!latitude!from!north!to! south.!!!!!!!!!!!!!!!!!!!!!!!
Figure*S5*(Page*43).!Mismatch!distributions!of!three!mitochondrial!haplogroups!observed!in! the!Micrurus(fulvius!complex!(solid!lines).!Simulated!distributions!under!a!model!of!spatial! expansion!are!shown!as!well!(dotted!lines).!Shapes!of!haplotypes!correspond!to!Fig.!S3.!
Figure*S6*(Page*44).!!Spearman’s!rho!correlation!coefficients!for!comparisons!between!latitude,! longitude!and!nuclear!DNA!spatial!principal!components!in!Micrurus(tener.!Spatial!principal! components!1–10!comprise!the!x@axis!and!demonstrate!that!higher!numbered!axes!have!a! Streicher et al. 2 weak!correlation!with!large@scale!spatial!structure.!Comparisons!were!performed!for!all! individuals!of!M.(tener!from!Mexico!and!the!United!States!(A!and!B),!and!individuals!of!M.(tener! collected!from!the!putative!range!front!in!the!United!States!(C!and!D).!All!correlation! coefficients!are!presented!as!absolute!in!order!to!display!the!magnitude!of!correlation!on!the! same!scale.!!
Figure*S7*(Page*45).!Lagged!scores!from!the!three!most!explanatory!nuclear!DNA!(nucDNA)! spatial!principal!components!(SPCs)!in!M.(tener!(A–C).!The!distribution!of!two!mitochondrial!
(mtDNA)!haplogroups!observed!across!the!same!individuals!used!in!the!SPCs!analysis!(D).!
*Figure*S8*(Page*46).!Alternative!species!distribution!models!for!last!glacial!maximum! predictions!using!the!MIROC@ESM!and!MPI@ESM@P!models.!!