ANNALS­­OF­­CARNEGIE­­MUSEUM voL. 80, numBer 4, pp. 323–350 31 auGuSt 2012

­­LITHOSTRATIGRAPHY,­PALEONTOLOGY,­BIOSTRATIGRAPHY, AND­AGE­OF­THE­UPPER­PALEOZOIC­ABO­FORMATION­NEAR­ JEMEZ­SPRINGS,­NORTHERN­NEW­MEXICO,­USA

Spencer G. LucaS Collections­and­Research­Department,­New­Mexico­Museum­of­Natural­History­and­Science,­ 1801­Mountain­Road­N.­W.,­Albuquerque,­New­Mexico­87104 [email protected]

SuSan K. HarriS Collections­and­Research­Department,­New­Mexico­Museum­of­Natural­History­and­Science,­ 1801­Mountain­Road­N.­W.,­Albuquerque,­New­Mexico­87104­ [email protected]

JuStin a. SpieLmann Collections­and­Research­Department,­New­Mexico­Museum­of­Natural­History­and­Science,­­ 1801­Mountain­Road­N.­W.,­Albuquerque,­New­Mexico­87104­ [email protected]

DaviD S Berman Curator,­Section­of­Vertebrate­Paleontology,­­ Carnegie­Museum­of­Natural­History,­Pittsburgh,­Pennsylvania­15213­ [email protected]

amy c. Henrici Collection­Manager,­Section­of­Vertebrate­Paleontology,­ ­Carnegie­Museum­of­Natural­History,­Pittsburgh,­Pennsylvania­15213­ [email protected]

KarL Krainer Institute­of­Geology­and­Paleontology,­ University­of­Innsbruck,­Innrain­52,­Innsbruck,­A-6020,­Austria­ [email protected]

Larry F. rineHart Collections­and­Research­Department,­New­Mexico­Museum­of­Natural­History­and­Science,­­ 1801­Mountain­Road­N.­W.,­Albuquerque,­New­Mexico­87104­ [email protected]

WiLLiam a. DimicHeLe Department­of­Paleobiology,­National­Museum­of­Natural­History,­ Smithsonian­Institution,­Washington,­D.C.­20560­ [email protected]

Dan S. cHaney Department­of­Paleobiology,­National­Museum­of­Natural­History,­­ Smithsonian­Institut ion,­Washington,­D.C.­20560­ [email protected]

HanS Kerp Forschungsstelle­für­Paläobotanik,­Insitut­für­Geologie­und­Paläontologie,­Westfälische­Wilhelms-Universität,­ Hindenburgplatz­57,­48143­­Münster,­Germany­ [email protected] 324 annaLS oF carneGie muSeum voL. 80

ABSTRACT

In­ the­ Jemez­ Springs­ area­ of­ Sandoval­ County,­ northern­ New­ Mexico,­ siliciclastic­ red­ beds­ of­ the­ upper­ Paleozoic­ Abo­ Formation­ are­ well­ exposed­and­yield­fossil­plants­and­vertebrates.­The­local­Abo­Formation­section­is­more­than­190­m­thick­and­rests­disconformably­on­the­Upper­ Pennsylvanian­Guadalupe­Box­Formation­and­is­conformably­overlain­by­the­Lower­Permian­DeChelly­Sandstone­(Yeso­Group).­Abo­sandstone­ sheets­are­low­sinuosity­river­deposits,­and­intercalated­sandstone­beds­and­lenses­represent­sheet­splays­and­minor­channel­fills­that­formed­during­ overbank­flooding.­The­dominant­Abo­lithofacies­is­mudstone,­which­represents­floodplain­deposits,­many­with­calcareous­paleosols.­Fossils­are­ present­in­three­stratigraphic­intervals­of­the­lower­to­middle­Abo­Formation.­All­three­intervals­yield­eupelycosaur-dominated­vertebrate­fossil­ assemblages­of­Coyotean­age­(Coyotean­=­late­Virgilian-Wolfcampian­on­the­North­American­provincial­marine­timescale:­Lucas­2006).­The­low- est­interval­also­yields­the­Spanish­Queen­Mine­paleoflora­of­pteridosperms­and­conifers.­Strata­of­the­Guadalupe­Box­Formation­disconformably­ below­the­Abo­Formation­contain­late­Virgilian­fusulinids.­We­correlate­the­Abo­Formation­fossil­assemblages­in­the­Jemez­Springs­area­to­the­ Coyotean-age­fossil­assemblages­in­the­upper­part­of­the­El­Cobre­Canyon­Formation­in­the­Arroyo­del­Agua­area­and­in­the­Cañon­del­Cobre­in­ the­Chama­basin­of­northern­New­Mexico.­This­suggests­a­middle­Wolfcampian­age­for­the­Jemez­Springs­area­fossil­assemblages,­an­age­very­ close­to­the­Pennsylvanian-Permian­boundary.

Key WorDS:­Coyotean,­Early­Permian,­paleoflora,­vertebrate­fossils,­Wolfcampian

INTRODUCTION USNM­ National­ Museum­ of­ Natural­ History, Smithsonian­Institution,­Washington,­D.C. In­northern­New­Mexico­the­Jemez­Mountains­are­an­im- WM­ Walker­Museum,­fossils­now­in­collection­of­ mense­late­Cenozoic­volcanic­edifice­located­on­the­west- the­Field­Museum­of­Natural­History,­Chicago,­ ern­edge­of­the­Rio­Grande­rift.­This­edifice­separates­the­ IL. southeastern­edge­of­the­San­Juan­Basin­to­the­west­and­ YPM Peabody­ Museum­ of­ Natural­ History,­ Yale­ southwest­from­the­Chama­River­basin­to­the­north­(Fig.­ University,­New­Haven,­CT. 1).­Along­the­southern­edge­of­the­Jemez­Mountains,­the­ Jemez­River­and­its­tributaries­are­cut­into­bedrock­can- yons­that­expose­sedimentary­rocks­as­old­as­Mississip- HISTORY­OF­FOSSIL­COLLECTING pian­ (e.g.,­ Wood­ and­ Northrop­ 1946;­ Woodward­ et­ al.­ Of­the­three­principal­Upper­Pennsylvanian-Lower­Perm- 1977;­Woodward­1987;­Osburn­et­al.­2002;­Kelley­et­al.­ ian­vertebrate­body­fossil­collecting­areas­in­north-central­ 2003).­ Particularly­ well-exposed­ outcrops­ of­ the­ Lower­ New­Mexico,­the­Jemez­Springs­area­(sometimes­referred­ Permian­Abo­Formation­are­present­along­the­Jemez­River­ to­as­the­“Cañon­[Canyon]­de­San­Diego”­locality)­has­re- south­of­Jemez­Springs­(Cañon­de­San­Diego)­and­along­ ceived­the­least­attention­from­vertebrate­paleontologists.­ the­ canyon­ of­ the­ Guadalupe­ River­ near­ Gilman.­ Fossil­ Whereas­ the­Arroyo­ del­Agua­ and­ Cañon­ del­ Cobre­ lo- vertebrates­were­discovered­in­these­Abo­Formation­red­ calities­(Fig.­1)­were­originally ­collected­in­the­late­1870s­ beds­during­the­1930s,­and­in­the­1950s­fossil­plants­were­ by­David­Baldwin,­discovery­of­vertebrate­fossils­in­the­ recognized­at­one­locality­south­of­Jemez­Springs.­During­ Jemez­Springs­area­was­not­made­until­more­than­half­a­ the­last­80­years,­the­paleontology­of­the­Abo­Formation­ century­later,­in­the­1930s. in­the­Jemez­Springs­area­has­been­intermittently­studied,­ ­ In­1931,­A.S.­Romer­of­Harvard­University­reasoned­ most­recently­by­field­crews­from­the­Carnegie ­Museum­of­ that­conditions­of­deposition­similar­to­those­of­the­nearby­ Natural­History­and­the­New­Mexico­Museum­of­Natural­ Arroyo­del­Agua­locality­might­exist­in­the­Abo­Formation­ History.­Here,­we­synthesize­the­results­of­this­research­to­ sediments­of­the­Jemez­Springs­area­(Fig.­2).­Consequent- provide­documentation­of­the­paleoflora­and­fossil­verte- ly,­he­prospected­the­walls­of­Cañon­de­San­Diego­and­met­ brate­assemblages­from­the­Abo­Formation­in­the­Jemez­ with­quick­success,­discovering­numerous­fossil­vertebrate­ Springs­area.­We­also­place­these­fossil­assemblages­into­ sites­that­yielded­bones­of­pelycosaurs­and­temnospondyls­ a­ detailed­ lithostratigraphic­ framework­ and­ interpret­ the­ (Romer­1960).­From­one­of­these­sites,­Romer­(1937)­de- age­and­correlation­of­the­Abo­Formation­in­the­vicinity­of­ scribed­the­well-preserved­partial­skeleton­of­a­new,­large­ Jemez­Springs. species­ of­ the­ eupelycosaur­ Sphenacodon Marsh,­ 1878,­ S. ferocior.­Seven­years­later,­unaware­of­Romer’s­brief,­ Institutional­Abbreviations but­productive­trip­to­the­Jemez­Springs­area,­a­field­party­ from­the­University­of­California­at­Berkeley­led­by­S.P.­ CM­ Carnegie­ Museum­ of­ Natural­ History,­ Pitts- Welles­also­prospected­the­walls­of­Cañon­de­San­Diego.­ burgh,­PA. They­collected­their­most­significant­find,­a­partial­embo- MCZ­ Museum­ of­ Comparative­ Zoology,­ Harvard lomerous­vertebral­column­described­by­Langston­(1953a)­ University,­Cambridge,­MA. as­ possibly­ pertaining­ to­ Archeria­ Case,­ 1918,­ from­ the­ NMMNH­ New­ Mexico­ Museum­ of­ Natural­ History,­ same­site­that­yielded­S. ferocior,­referred­to­as­the­Span- Albuquerque,­NM. ish­Queen­Mine­locality­(Berman­1993).­Approximately­ UCMP University­ of­ California­ Museum­ of­ Paleon- 3­km­north­of­this­locality,­and­90­m­north­of­the­copper­ tology,­Berkeley,­CA. mine­ itself,­ the­ University­ of­ California­ field­ party­ also­ USGS U.S.­Geological­Survey,­Washington,­D.C. discovered­a­fossil­vertebrate­site­stratigraphically­low­in­ 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­325 the­Abo­section,­which­they­named­the­Johnson­locality,­ that­yielded­bones­referable­to­small­diadectids­and­“sphe- nacodonts”­ (Langston­ 1953a,­ 1953b;­ Berman­ and­ Reisz­ 1980;­Berman­1993;­Eberth­and­Berman­1993). ­ A­hiatus­of­nearly­four­decades­ensued­before­collect- ing­in­the­Jemez­Springs­area­began­anew­with­field­par- ties­from­the­Carnegie­Museum­of­Natural­History­under­ the­ direction­ of­ D.­ S­ Berman­ from­ 1975­ to­ 1978.­ Like­ Romer,­the­Carnegie­Museum­parties­discovered­multiple­ vertebrate­fossil­localities­at­various­intervals­in­the­lower­ part­of­the­Abo­Formation­section.­Their­discovery­of­a­ partial,­articulated­sail­and­lower­jaw­elements­of­a­new,­ small­species­of­Dimetrodon­Cope,­1878,­D. occidentalis,­ marked­the­first­recorded­co-occurrence­of­the­eupelyco- saur­genera­Dimetrodon­and­Sphenacodon (Berman­1977).­ Another­site,­just­north­of­the­confluence­of­the­Guadalupe­ and­Jemez­rivers,­yielded­a­nearly­complete­skull­and­post- crania­ described­ by­ Berman­ and­ Reisz­ (1980)­ as­ a­ new­ species­of­Trimerorhachis­Cope,­1878,­T. sandovalensis.­ Additional­finds­of­mostly­isolated,­though­diagnostic,­el- ements­pertaining­to­Xenacanthus Beyrich,­1848,­Eryops­ Cope,­1877,­Zatrachys­Cope,­1878,­Platyhystrix­Williston,­ 1911,­Diplocaulus­Cope,­1877,­and­Diadectes­Cope,­1878,­ more­than­doubled­the­number­of­vertebrate­taxa­known­ from­ the­ Jemez­ Springs­ area­ (Berman­ and­ Reisz­ 1980;­ Berman­1993). ­ From­2003­through­2007,­joint­parties­from­the­Carn- egie­ Museum­ of­ Natural­ History­ and­ New­ Mexico­ Mu- seum­ of­ Natural­ History­ revisited­ the­ known­ localities­ mentioned­above­in­order­to­establish­their­stratigraphic­ context­ within­ the­ Abo­ Formation­ section­ near­ Jemez­ Springs.­During­the­last­decade,­New­Mexico­Museum­of­ Natural­History­field­crews­discovered­several­new­sites­ from­ which­ small­ quantities­ of­ surface-collected­ fossils­ were­recovered.­At­present,­we­are­aware­of­18­discrete­ vertebrate­fossil­localities­in­the­Abo­Formation­in­the­Je- mez­Springs­area­that­we­can­place­into­a­detailed­strati- graphic­context­(Fig.­2). ­ During­ the­ 1950s,­ Sergius­ Mamay­ and­ Ellis­ Yochel- son,­both­of­the­U.S.­Geological­Survey,­collected­a­small­ paleoflora­ at­ the­ Spanish­ Queen­ Mine,­ south­ of­ Jemez­ Springs­(Fig.­2).

Fig­1.—Index­map­of­northern­New­Mexico­showing­location­of­prin- GEOLOGY­AND­STRATIGRAPHIC­CONTEXT cipal­ Pennsylvanian-Permian­ fossil­ collecting­ areas­ in­ Chama­ River­ basin­(Cañon­del­Cobre,­Arroyo­del­Agua)­and­in­Jemez­Springs­area.­ In­ the­ Jemez­ Springs­ area­ the­Abo­ Formation­ is­ exten- Modified­from­Woodward­(1974). sively­exposed­along­canyon­walls,­terraces,­and­tributary­ arroyos­of­the­Jemez­and­Guadalupe­rivers­(Fig.­3).­We­ The­Abo­red­beds­along­the­Jemez­River­are­overlain­by­ measured­eight­stratigraphic­sections­in­the­Abo­Forma- eolian­sandstone­of­the­DeChelly­Sandstone­of­the­Yeso­ tion­in­the­Jemez­Springs­area­to­establish­the­stratigraphic­ Group­ (formerly­ called­ the­ “Meseta­ Blanca­ Sandstone­ distribution­ of­ the­ 18­ vertebrate­ fossil­ localities­ and­ the­ Member­ of­ the­ Yeso­ Formation:”­ Lucas­ et­ al.­ 2005a).­ one­fossil­plant­locality­that­we­can­locate­in­the­local­Abo­ Along­the­Guadalupe­River­at­Gilman,­the­Abo­Formation­ section­(Fig.­4;­Appendices­1–2).­ is­completely­exposed.­Here­the­measured­thickness­is­131­ ­ Along­the­Jemez­River,­the­maximum­measured­thick- m,­and­the­Abo­Formation­is­underlain­by­the­Upper­Penn- ness­of­the­Abo­Formation­is­190­m,­but­the­base­of­the­ sylvanian­Guadalupe­Box­Formation­(Krainer­et­al.­2005)­ Abo­ Formation­ is­ not­ exposed­ south­ of­ Jemez­ Springs.­ and­overlain­by­a­few­meters­of­Yeso­Group­red­beds­and­ 326 annaLS oF carneGie muSeum voL. 80

Fig.­2.—Map­of­the­Jemez­Springs­area­showing­fossil­localities­and­locations­of­measured­stratigraphic­sections­in­the­Abo­Formation.­Precise­map­ coordinates­for­all­fossil­localities­are­on­file­at­the­CM,­NMMNH,­and­USNM. 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­327

Fig.­3.—Photographs­of­selected­Abo­Formation­outcrops­in­the­Jemez­Springs­area.­A,­view­looking­west­from­Dimetrodon­site­section.­The­saddle­ on­the­skyline­is­between­Mesa­de­Guadalupe­(on­the­left)­and­Virgin­Mesa­(on­the­right).­Both­mesas­are­capped­by­Pleistocene­Bandelier­Tuff­overly- ing­cliff-­and­ledge-forming­sandstone­beds­of­the­Lower­Permian­DeChelly­Sandstone­(Yeso­Group)­above­slopes­of­Abo­Formation.­B, characteristic­ sheet­sandstone­of­Abo­Formation­in­Dimetrodon­site­section.­C,­calcrete­nodules­and­rhizoliths­in­Abo­Formation­mudstone.­D,­trough­crossbedded,­ pebbly­sandstone­of­Abo­Formation. lower­Pleistocene­Bandelier­Tuff­(Fig.­4). at­least­20­m­above­the­local­base­of­the­Abo­Formation.­ ­ Throughout­the­Jemez­Springs­area,­the­Abo­Formation­ It­ encompasses­ the­ UCMP­ Johnson­ locality,­ the­ USNM­ can­be­divided­into­lower­and­upper­members­(Fig.­4).­The­ Spanish­ Queen­ Mine­ locality,­ and­ NMMNH­ Locality­ lower­member­is­dominated­by­mudstone­and­siltstone­that­ 7661. constitute­69-72%­of­the­Abo­section.­In­the­upper­mem- ­ 2.­Assemblage­B­consists­of­two­NMMNH­vertebrate­ ber,­mudstone­and­siltstone­constitute­much­less­(25-41%)­ fossil­localities­(4507­and­5381)­at­least­40­m­above­the­ of­the­Abo­section.­The­thickness­of­the­lower­member­is­ base­of­the­local­Abo­section­(Fig.­4). variable,­at­least­115­m­thick­along­the­Jemez­River­and­ ­ 3.­Assemblage­C­encompasses­most­of­the­Abo­Forma- about­90­m­thick­at­Gilman,­whereas­the­upper­member­is­ tion­vertebrate­fossil­localities­in­the­Jemez­Springs­area­ more­uniform­and­about­60­m­thick­along­the­Jemez­River­ (CM­ localities:­ 999,­ 1000­ [=­ NMMNH­ Locality­ 7056],­ and­at­Gilman.­These­members­probably­are­homotaxial­to­ 1001­[=­NMMNH­Locality­5380],­1002­[=­NMMNH­Lo- the­Scholle­Member­(lower)­and­Cañon­de­Espinoso­Mem- cality­7055],­1003,­4047,­4048,­4049,­4050;­NMMNH­lo- ber­(upper)­of­the­Abo­type­section­in­central­New­Mexico­ calities:­4035,­4508,­5366,­7057,­7060,­7061,­7062;­YPM­ (Lucas­et­al.­2005a). lungfish­locality)­and­is­from­an­~30-m-thick­interval­that­ ­ Fossil­ localities­ in­ the­ Abo­ Formation­ in­ the­ Jemez­ begins­about­90­m­above­the­base­of­the­local­Abo­section­ Springs­area­can­be­assigned­to­three­stratigraphic­inter- (Fig.­4).­In­other­words,­it­is­in­about­the­middle­of­the­ vals­(Fig.­4): local­Abo­section­and­straddles­the­boundary­between­the­ ­ 1.­ Assemblage­ A­ is­ the­ lowest­ stratigraphic­ interval,­ lower­and­upper­members­of­the­Abo­Formation. 328 annaLS oF carneGie muSeum voL. 80

Fig.­4.—Measured­stratigraphic­sections­of­the­Abo­Formation­in­the­Jemez­Springs­area­showing­stratigraphic­distribution­of­fossil­localities.­See­ Appendix­1­for­map­coordinates­of­the­measured­sections­and­Appendix­2­for­a­list­of­fossil­localities­and­taxa. 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­329

­ Romer’s­(1937,­1960)­term­“Spanish­Queen­Mine­lo- ward­decrease­in­bed­thickness­and­grain­size.­Common­ cality”­does­not­indicate­the­same­stratigraphic­level­as­the­ lithofacies­are­clast-supported­conglomerates­at­the­base,­ actual­mine,­which­is­the­site­of­the­fossil­plant­collection.­ grading­into­multistoried,­trough-crossbedded­sandstone,­ Romer­used­the­nearest­distinctive­geographic­place­name­ which­in­the­upper­part­may­be­associated­with­horizontal- for­the­collections­he­made,­which,­in­this­instance,­were­ ly­laminated,­massive,­low-angle­crossbedded­or­trough- not­right­at­the­mine­(Langston­1953b;­Berman­1993;­Hunt­ crossbedded­sandstone­(e.g.,­Fig.­3D).­Sometimes­in­the­ and­Lucas­1996).­Indeed,­Langston­(1953b:­fig.­4)­located­ uppermost­part­of­the­fine-grained­sandstone,­the­strata­are­ the­Spanish­Queen­Mine­(vertebrate)­locality­(presumably­ bioturbated,­ and­ synsedimentary­ deformation­ structures­ based­on­information­provided­to­him­by­Romer)­about­3­ are­rarely­observed.­Sandstone­ribbons­have­not­been­ob- km­south-southwest­of­the­Johnson­locality­in­the­vicinity­ served­at­Jemez­Springs­Abo­Formation­outcrops,­suggest- of­our­Dimetrodon­site­section­(Fig.­2).­As­this­is­the­most­ ing­that­such­channelized­deposits­were­rare­in­this­area­ fossiliferous­Abo­outcrop­area,­with­most­of­the­localities­ (Eberth­and­Miall­1991;­Eberth­and­Berman­1993). relatively­high­stratigraphically,­in­our­assemblage­C,­we­ ­ The­abundance­of­internal­scours­and­erosional­surfac- conclude­ that­ Romer’s­ collections­ came­ from­ the­ strati- es,­presence­of ­conglomeratic­lags­at­the­base,­variation­in­ graphic­level­of­assemblage­C.­ grain­ size,­ and­ poorly-­ to­ well-developed­ fining-upward­ ­ Note­that­assemblages­A­and­B­encompass­only­a­few­ sequences­ are­ indicative­ of­ ephemeral­ fluvial­ regimes­ localities­ over­ a­ short­ stratigraphic­ interval­ (Fig.­ 4).­ So,­ (e.g.,­Picard­and­High­1973;­Tunbridge­1981,­1984;­Mi- they­ could­ be­ combined­ to­ form­ one­ assemblage.­ How- all­1996).­Sheet­sandstones­are­interpreted­as­deposits­of­ ever,­we­recognize­three­stratigraphically-separated­fossil­ broad,­shallow­channels­of­a­probable­braided­river­sys- assemblages­in­the­Abo­Formation­in­the­Jemez­Springs­ tem.­The­lack­of­lateral­accretion­deposits­indicates­that­ area,­simply­because­this­is­the­most­precise­way ­to­depict­ the­channels­were­shallow,­so­that­large­macroform­bars­ the­distribution­of­fossils­in­the­section.­ could­not­develop.­The­sandstone­sheets­are­very­similar­ ­ Looff­(1987),­Eberth­and­Miall­(1991),­Eberth­and­Ber- to­the­“major­sandstone­sheets”­of­the­Cutler­Group­de- man­ (1993),­ and­ Krainer­ and­ Lucas­ (2010)­ studied­ the­ scribed­in­detail­by­Eberth­and­Miall­(1991).­These­authors­ sedimentology­of­the­Abo­Formation­in­the­Jemez­Springs­ compared­ the­ sheet­ sandstones­ to­ those­ of­ the­ modern­ area.­Krainer­and­Lucas­(2010)­concluded­that­sandstone­ Platte­River­(model­9­of­Miall­1985). sheets­in­the­Abo­were­deposited­by­low­sinuosity­river­ ­ In­the­Abo­Formation­near­Jemez­Springs, thin­inter- channels.­More­lenticular­sandstone­beds­represent­minor­ calated­sandstone­beds­and­lenses­are­mostly­10­to­30­cm,­ channel­fills­and­crevasse­splay­deposits.­Most­of­the­Abo­ rarely­up­to­50­cm­thick.­They­occur­as­single­sandstone­ Formation­in­the­Jemez­Springs­area­consists­of­mudstone.­ beds­that­are­laminated,­massive,­or­crossbedded.­Stacked­ Many­mudstone­beds­contain­calcrete­nodules­and­rhizo- sandstone­units­are­up­to­1­m­thick.­The­sandstone­beds­ liths­(e.g.,­Fig.­3C)­indicative­of­the­development­of­cal- occur­ as­ tabular­ or­ lens-shaped­ bodies;­ their­ bases­ may­ careous­ paleosols.­ The­ mudstone­ lithofacies­ of­ the­Abo­ be­erosive.­Rarely,­sandstone­sheets­are­bioturbated.­This­ Formation­is­interpreted­as­floodplain­deposits.­No­pond­ architectural­ element­ is­ similar­ to­ the­ “minor­ sandstone­ or­lake­deposits­have­been­identified­in­the­Abo­Formation­ sheets­and­lenses”­that­Eberth­and­Miall­(1991)­described­ in­the­Jemez­Springs­area. from­the­Cutler­Group­red ­beds­in­the­Chama­basin­to­the­ ­ According­ to­ Miall­ (1985,­ 1992,­ 1996),­ architectural­ north. elements­ of­ depositional­ systems­ are­ characterized­ by­ a­ ­ Intercalated,­ laterally-restricted­ sheet­ sands­ and­ len- distinctive­facies­assemblage,­internal­geometry,­external­ ses­ are­ characteristic­ of­ overbank­ flooding,­ particularly­ form­and,­partly,­by­a­distinct­vertical­profile.­Each­type­ in­ ephemeral­ systems­ (e.g.,­ Williams­ 1971;­ Picard­ and­ represents­ a­ particular­ process­ occurring­ within­ a­ depo- High­1973;­Tunbridge­1981).­The­tabular­sandstone­beds­ sitional­ system.­Within­ the­Abo­ Formation,­ Krainer­ and­ are­ probably­ sheet­ splays.­ Basal­ erosional­ surfaces­ with­ Lucas­(2010)­identified­three­architectural­elements–sand- mudstone­ rip-up­ clasts­ and­ reworked­ pedogenic­ carbon- stone­sheets,­intercalated­sandstone­beds­and­lenses,­and­ ate­clasts­indicate­high-energy­transport.­Lack­of­internal­ siltstone-mudstone. erosion­ and­ reactivation­ surfaces­ within­ the­ intercalated­ ­ The­most­characteristic­and­distinct­facies­assemblages­ sandstone­beds­is­typical­of­sheet­flow.­Sandstone­lenses­ in­the­Abo­Formation­are­sandstone­sheets­(e.g.,­Fig.­3B),­ represent­minor­channel­fills­and­may­represent­the­feeder­ which­correspond­to­the­architectural­element­CH­(chan- channels­(crevasse­channels)­of­the­sheet­splays. nel).­Sandstone­sheets­are­characterized­by­width-to-depth­ ­ Siltstone­and­mudstone­units­are­the­dominant­lithofa- ratios­ greater­ than­ 15:1,­ whereas­ sandstone­ ribbons­ also­ cies­of­the­Abo­Formation­in­the­Jemez­Springs­area,­oc- corresponding­to­the­element­CH,­have­ratios­smaller­than­ curring­as­massive­or­laminated­beds,­some­with­dessica- 15:1.­Within­the­Abo­Formation­in­the­Jemez­Springs­area,­ tion­cracks­and/or­rhizoliths.­A­few­pedogenic­carbonate­ sandstone­sheets­form­prominent,­resistant­ledges­that­can­ nodular­zones­are­present.­The­siltstone-mudstone­facies­ be­ traced­ laterally­ over­ long­ distances­ (commonly­ more­ that­forms­sheet-like­units­extending­laterally­over­at­least­ than­100­m,­up­to­several­hundred­meters).­The­base­of­ hundreds­of­meters­belongs­to­element­FF­(floodplain­fine)­ any­given­sandstone­body­is­generally­erosive.­Sublithofa- of­ Miall­ (1996)­ and­ is­ interpreted­ as­ floodplain­ depos- cies­within­the­sandstone­bodies­commonly­show­an­up- its­formed­by­settling­from­sheetfloods.­The­presence­of­ 330 annaLS oF carneGie muSeum voL. 80

Fig.­5.—Selected­fossil­plants­from­the­Spanish­Queen­Mine­paleoflora.­A,­Autunia naumanii,­USNM­539289,­nearly­complete­pinna.­B,­Walchia piniformis, USNM­539299,­branch­fragment.­C,­Neurodontopteris auriculata,­USNM­538957,­pinna­fragment.­D,­Ernestiodendron filiciforme,­USNM­ 539292,­branch­fragment.­E,­Rhachiphyllum schenkii,­USNM­539314,­frond­segment­with­several­partial­pinnae.­Note­flatness­of­pinnule­laminae,­ which­contrasts­with­Autunia conferta,­wherein­the­laminae­are­vaulted­and­much­more­robust.­F,­Taeniopteris­sp.,­USNM­539308,­one-half­of­lamina­ showing­midrib­on­right­side.­Scale­bars­=­1­cm. 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­331

Fig.­6.—Xenacanthus,­Gnathorhiza, Eryops,­Diplocaulus, and­Platyhystrix­from­the­Abo­Formation­in­the­Jemez­Springs­area.­A-C, Eryops sp.,­CM­ 26567,­A-B,­neural­spine­in­lateral­views­and­C,­intercentrum,­from­NMMNH­Locality­7056.­D-I, Platyhystrix sp.,­CM­38029,­neural­spine­fragments,­ from­CM­Locality­5,­in­lateral­view.­J,­Xenacanthus­sp.,­CM­38026,­tooth­base,­from­CM­Locality­1,­in­apical­view.­K,­Gnathorhiza sp.,­YPM­8636,­tooth­ plate,­from­“east­of­highway­4­and­south­of­Jemez­Springs,”­in­occlusal­view­(photograph­courtesy­of­Chris­Norris).­L, Diadectidae,­NMMNH­P-51876,­ partial­maxilla,­from­NMMNH­Locality­6842,­in­lateral­view.­M-N,­Diplocaulus sp.,­NMMNH­P-61686,­vertebra,­from­NMMNH­Locality­6842,­in­ M, anterior­and­N,­lateral­views.­Scale­bars:­A-B, upper­left;­C-E,­upper­right;­F-I,­middle­left;­J-K,­middle­right;­L,­lower­left;­and­M-N,­lower­right. 332 annaLS oF carneGie muSeum voL. 80

River­flows­(Fig.­2).­Fossil­plants­collected­from­this­mine­ Table 1. Plant­taxa­from­the­Spanish­Queen­Mine­paleoflora. (Fig.­5)­by­Mamay­and­Yochelson­are­in­the­USNM­collec- Sphenopsids: tion­and­have­not­previously­been­documented.­They­occur­ ­ Annularia spicata in­light­to­medium­gray,­micaceous­sediment­that­grades­ ­ Calamostachys spicata­ from­siltstone­to­fine­sandstone.­Plant­fossil­remains­range­ Ferns: from­large­portions­of­fronds­to­organic­hash,­the­latter­be- Asterotheca­sp. ­ Polymorphopteris­sp. ing­abundant.­The­siltstone­contains­mm-scale­laminations­ ­ Pecopteris bredovii with­occasional­thicker­lenses.­Organic­remains­are­found­ Pteridosperms: on­ most­ bedding­ surfaces.­ Preservational­ detail­ is­ good,­ ­ Alethopteris schneideri but­the­cuticle­appears­to­be­preserved­only­on­the­most­ ­ Callipteridium gigas robust­remains,­such­as­conifer­foliage. ­ Neurodontopteris auriculata ­ The­ late­ S.­ Mamay­ (personal­ communication­ to­ W.­ Callipterids: DiMichele­1995)­reported­that­the­fossil­plant­collection­ ­ Autunia naumanii ­ Dichophyllum flabellifera from­ the­ Spanish­ Queen­ Mine­ came­ from­ inside­ one­ of­ Dichophyllum moorei the­larger­mine­adits­and­was­taken­either­directly­from­ Rhachiphyllum schenkii the­roof­or­from­roof­falls­not­far­from­the­adit­entrance.­ Conifers: Over­the­intervening­years,­the­adits­have­become­inac- ­ Culmitzschia/Lebachia speciosa cessible,­making­it­impossible­to­relocate­the­exact­spot­ ­ Ernestiodendron filiciforme from­ which­ the­ fossil­ plant­ collection­ was­ made.­ How- Walchia piniformis ­ Gomphostrobus bifidus ever,­fossil-plant-bearing­siltstones­of­the­same­character­ as­those­collected­by­Mamay­and­Yochelson­are­present­ Incertae­sedis: ­ Taeniopteris­sp. in­the­strata­observable­in­the­rock­faces­between­the­old­ adits.­Thus,­ the­ fossil-plant-bearing­ beds­ occur­ within­ a­ widespread­sandstone­body,­up­to­4­m­thick,­consisting­of­ lamination­ indicates­ deposition­ from­ suspension.­ How- scours­and­lenses­of­finely­laminated­micaceous­siltstone.­ ever,­ stratification­ is­ mostly­ absent,­ probably­ due­ to­ the­ The­organic-rich­laminated­beds­are­tens­of­cms­thick­and­ presence­ of­ organisms­ (bioturbation­ or­ pedoturbation).­ appear­to­represent­bars­within­a­braided­stream­channel­ Desiccation­cracks,­particularly­in­the­finer-grained­facies,­ system.­The­large­amount­of­highly­comminuted­plant­re- indicate­ periodic­ drying.­A­ strong­ seasonal­ moisture­ re- mains­indicates­some­degree­of­destruction­of­plant­mate- gime­with­a­period­of­high­evapo-transpiration­facilitated­ rial­either­by­grinding­in­bed­load­or­by­the­sweeping­of­ the­formation­of­calcic­paleosols. partially­decayed­material­by­runoff­from­the­paleosol­sur- ­ The­average­petrographic­composition­of­the­Abo­For- faces­surrounding­the­active­channel­(e.g.,­Gastaldo­1991). mation­at­the­Jemez­River­and­Gilman­is­very­similar­to­ ­ The­elements­of­the­paleoflora­are­summarized­in­Ta- that­of­the­Cutler­Group­at­Cañon­del­Cobre­and­Arroyo­ ble­1.­The­most­abundant­of­these­elements,­based­on­an­ del­Agua­in­the­Chama­basin­to­the­north­(Krainer­and­Lu- assessment­of­this­small­collection,­are­the­pteridosperm­ cas­2010).­Sandstones­are­dominantly­arkoses­and­lithic­ arenites,­subordinately­subarkoses.­The­main­source­rock­ Neurodontopteris auriculata­ Potonié,­ 1893,­ callipterids,­ type­ is­ granite,­ from­ which­ the­ majority­ of­ mono-­ and­ particularly­Autunia naumanii (Gutbier)­Schimper,­1869,­ polycrystalline­quartz,­almost­all­detrital­feldspars­and­mi- and­conifers,­particularly­Culmitzschia/Lebachia speciosa­ cas,­and­all­granitic­rock­fragments­were­derived.­Meta- (Florin)­Clenent-Westerhof,­1984.­Several­taxa­are­present­ morphic­rocks­provided­a­few­mono-­and­polycrystalline­ in­low­numbers­but­are­common­enough­to­be­present­con- quartz,­and­metamorphic­rock­fragments.­Most­sedimen- sistently­throughout­the­collectio n,­including­Rhachiphyl- tary­rock­fragments­were­derived­from­the­reworking­of­ lum schenkii (Heyer)­Kerp,­1988,­Ernestiodendron filici- pedogenic­ carbonate;­ subordinately,­ reworked­ mudstone­ forme­ (Sternberg)­ Florin,­ 1934,­ and­ laminae­ of­ several­ and­siltstone­grains­are­present.­Sandstone­is­commonly­ different­ sizes of­ Taeniopteris­ Brongniart,­ 1828.­ Lower­ cemented­ by­ coarse,­ blocky­ calcite;­ quartz­ cement­ may­ vascular­plants,­the­sphenopsids­and­ferns,­are­exceedingly­ also­be­present. rare,­ represented­ only­ by­ small­ numbers­ of­ fragmentary­ specimens.­ PALEOBOTANY-FLORA­OF­THE­SPANISH­QUEEN­ MINE­LOCALITY VERTEBRATE­PALEONTOLOGY The­Spanish­Queen­Mine­fossil­plant­locality­(USGS­Lo- We­provide­a­comprehensive­review­of­the­vertebrate­fos- cality­8936­and­USNM­Locality­43566)­is­at­a­long-closed­ sils­collected­from­the­Abo­Formation­in­the­Jemez­Springs­ copper­ mine­ located­ approximately­ 2­ km­ southwest­ of­ area.­Taxa­discussed­include­a­chondrichthyan,­a­dipnoan,­ Jemez­ Springs­ (Read­ and­ Mamay­ 1964),­ on­ the­ eastern­ four­ temnospondyl­ amphibian­ taxa,­ an­ anthracosaur,­ a­ side­of­the­Cañon­de­San­Diego,­through­which­the­Jemez­ nectridean,­a­diadectomorph,­and­four­eupelycosaur­taxa. 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­333

Fig.­ 7.—UCMP­ Johnson­ locality­ (UCMP­ Locality­ V3740)­ specimens­ from­ the­ Abo­ Formation­ of­ the­ Jemez­ Springs­ area.­ A-B,­ UCMP­ 195504,­ Sphenacodontidae­indet.,­right­femur­in­A,­dorsal­and­B,­ventral­views.­C-H,­Ophiacodon sp.,­three­dorsal­vertebrae­in­C (UCMP­195505), E (UCMP­ 195506), G (UCMP­195507),­anterior­and­D (UCMP­195505), F (UCMP­195506), H (UCMP­195507),­posterior­views.­I-K,­Eryops sp.,­I,­UCMP­ 195508,­ornamented­dermal­bone,­J,­UCMP­195509,­partial­palate­with­teeth,­K,­UCMP­195510,­premaxilla­with­two­large­teeth.­L,­UCMP­195511,­ Diadectidae­indet.,­right­mandible­in­lateral­view.­Photographs­courtesy­of­P.­Holroyd. 334 annaLS oF carneGie muSeum voL. 80

SYSTEMATIC­PALEONTOLOGY Eryops­ (Berman­ and­ Reisz­ 1980).­ Further,­ Berman­ and­ Class­Chondrichthyes­Huxley,­1880 Reisz­noted­the­probable­assignment­of­isolated­rhachito- Order­Elasmobranchii­Bonaparte,­1838 mous­vertebrae­collected­in­1938­from­the­Johnson­local- Xenacanthus­Beyrich,­1848­ ity­by­the­University­of­California­(Langston­1953b)­to­the­ genus­Eryops.­Additional­temnospondyl­amphibian­mate- Xenacanthus sp. rial­recovered­from­the­Johnson­locality­(UCMP­Locality­ (Fig.­6J) V3740)­in­the­UCMP­collection­that­we­assign­to­Eryops­ includes­a­partial­palate­with­teeth­(Fig.­7J),­a­premaxilla­ Referred specimen.—From­CM­Locality­999:­CM­38026,­ with­two­large­teeth­(Fig.­7K),­and­dermal­bone­with­pit- tooth­(Fig.­6J).­ and-ridge­ornamentation­(Fig.­7I).­ Discussion.—A­single,­incomplete­tooth­with­both­crowns­ ­ Three­ additional­ sites­ (NMMNH­ L-5381,­ 4507,­ and­ broken­off­is­referable­to­the­freshwater­shark­Xenacan- 7062)­in­Cañon­de­San­Diego­have­yielded­single,­isolated­ thus­(Berman­and­Reisz­1980;­Berman­1993).­ rhachitomous­ vertebral­ elements­ possibly­ pertaining­ to­ Eryops­(Figs.­8A–F).­Langston­(1953b)­collected­a­small,­ poorly­preserved­amphibian­skull­(UCMP­39179)­from­the­ Class­Osteichthyes­Huxley,­1880 Spanish­Queen­Mine­vertebrate­locality­(UCMP­V3855)­ Order­Dipnoi­Müller,­1844 in­the­same­matrix­block­from­which­he­recovered­articu- Gnathorhiza Cope,­1883 lated,­embolomerous­vertebrae­possibly­pertaining­to­Ar- Gnathorhiza sp.­ cheria­(see­below).­Although­Langston­(1953b:377)­noted­ (Fig.­6K) the­ skull­ has­ “no­ particularly­ diagnostic­ characters,”­ he­ provisionally­assigned­it­to­Eryops grandis­(Marsh,­1878),­ Referred specimens. —From­a­locality­east­of­the­high- based­on­the­marginal­and­palatal­teeth. way­and­south­of­Jemez­Springs­in­Cañon­de­San­Diego:­ YPM­8636­(Fig.­6K),­8637,­tooth­plates.­ Family­Trimerorhachidae­Cope,­1878 Discussion.—Two­tooth­plates­of­the­aestivating­lungfish­ Trimerorhachis­Cope,­1878 Gnathorhiza­were­collected­from­a­locality­in­Cañon­de­San­ Trimerorhachis sandovalensis­Berman­and­Reisz,­1980 Diego­by­M.­Fraccasso­and­S.G.­Lucas­while­students­at­ Yale­University­(Berman­and­Reisz­1980;­Berman­1993). Referred specimens.—­ From­ CM­ Locality­ 1001:­ CM­ 38027,­partial­skull­with­postcranial­elements.­From­CM­ Class­Amphibia­Linnaeus,­1758 Locality­ 1002:­ CM­ 38025,­ holotype,­ partial­ articulated­ Order­Temnospondyli­Zittel,­1888 skeleton­(Berman­and­Reisz­1980:­figs.­1–8).­ Family­Eryopidae­Cope,­1882 Discussion.—Berman­and­Reisz­(1980)­described­a­new­ Eryops Cope,­1877 trimerorhachid­ species,­ Trimerorhachis­ sandovalensis,­ Eryops sp. based­on­a­nearly­complete­skull­and­large­portion­of­an­ (Figs.­6A–C,­7I–K,­8A–F) articulated­ postcranial­ skeleton.­ The­ specimen­ was­ pre- served­ on­ three­ separate­ matrix­ blocks­ collected­ from­ a­ Referred specimens.—­ From­ CM­ Locality­ 1000:­ CM­ thick­sequence­of­light­red­mudstones­directly­surrounded­ 26567,­ vertebral­ intercentrum­ and­ neural­ spine­ (Figs.­ by­ poorly­ consolidated,­ coarse-grained,­ light­ gray­ sedi- 6A–C)­and­CM­86177,­jaw­fragments.­From­CM­Locality­ ments­of­a­very­small­stream­channel­lens. 4048:­CM­86170,­neural­spine.­From­NMMNH­Locality­ 4507:­ NMMNH­ P-32230,­ vertebral­ intercentrum­ (Figs.­ Superfamily­Dissorophoidea­Bolt,­1969 8C–D).­From­NMMNH­Locality­5381:­NMMNH­P-61641,­ Family­Dissorophidae­Boulenger,­1902 vertebral­intercentrum­(Figs.­8A–B).­From­NMMNH­Lo- Platyhystrix Williston,­1911 cality­7062:­NMMNH­P-61672,­vertebral­pleuracentrum­ (Figs.­8E–F).­From­the­Johnson­locality,­UCMP­Locality­ Platyhystrix­sp. V3740:­ UCMP­ 195508,­ ornamented­ dermal­ bone­ (Fig.­ (Figs.­6D–I) 7I);­UCMP­195509,­partial­palate­with­teeth­(Fig.­7J);­and­ UCMP­195510,­premaxilla­with­two­large­teeth­(Fig.­7K).­ Referred specimens.—From­ CM­ Locality­ 1003:­ CM­ From­Spanish­Queen­Mine­locality,­UCMP­V3855:­UCMP­ 38029,­neural­spine­fragments­(Figs.­6D–I). 39179,­small,­fragmentary­skull. Discussion.—Several­neural­spine­fragments­of­the­disso- Discussion.—Prior­ to­ this­ article,­ only­ one­ specimen,­ rophid­Platyhystrix­were­recovered­from­a­single ­locality­ a­ single­ vertebral­ intercentrum­ and­ neural­ spine­ (Figs.­ in­the­Jemez­Springs­area.­The­distinctive,­pustulose­pat- 6A–C),­from­CM­Locality­1000,­had­been­identified­with­ terning­along­the­neural­spines­identifies­them­as­Platyhys- confidence­ as­ pertaining­ to­ the­ semi-terrestrial­ eryopid­ trix­(Berman­and­Reisz­1980;­Berman­1993).­ 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­335

Fig.­8.—­Amphibia­and­Sphenacodontidae­from­the­Abo­Formation­in­the­Jemez­Springs­area.­A-F,­Amphibia.­A-B,­NMMNH­P-61641,­Eryops sp.,­ intercentrum,­from­NMMNH­Locality­5381,­in­A,­anterior/posterior­and­B,­ventral­views.­C-D,­NMMNH­P-32230,­Temnospondyli,­rhachitomous­ intercentrum,­from­NMMNH­Locality­4507,­in­C,­anterior/posterior­and­D,­ventral­views.­E-F,­NMMNH­P-61672,­Eryops­sp.,­pleurocentrum,­from­ NMMNH­ Locality­ 7062,­ in­ E,­ anterior/posterior­ and­ F, ventral­ views.­ G-O,­ Dimetrodon­ sp.,­ G-I,­ NMMNH­ P-61635,­ three­ neural­ spines,­ from­ NMMNH­Locality­5366,­in­G-H,­J-K,­M-N,­lateral­and­I,­L,­O,­cross­sectional­views.

Family­Zatrachyidae­Cope,­1878 sculpturing­clearly­identifies­it­(Berman­1993).­ Zatrachys Cope­1878 Zatrachys sp. Order­Anthracosauria­Säve-Söderbergh,­1937 Family­Archeriidae­Kuhn,­1965 Referred specimen.—From­ CM­ Locality­ 1000:­ CM­ Archeria Case,­1918 41706,­tabular. ?Archeria sp. Discussion.—The­presence­of­Zatrachys­is­known­in­the­ Jemez­ Springs­ vertebrate­ assemblage­ based­ on­ a­ single­ Referred specimens.—From­the­Spanish­Queen­Mine­ver- well-preserved­tabular­bone­whose­distinctive­shape­and­ tebrate­locality,­UCMP­V3855:­UCMP­39179,­articulated­ 336 annaLS oF carneGie muSeum voL. 80

Fig.­9.—Sphenacodontidae­and­Diadectidae­from­the­Abo­Formation­in­the­Jemez­Springs­area.­A-I.­Sphenacodontidae.­A-B,­NMMNH­P-32236,­ Sphenacodontidae,­ vertebral­ centrum­ and­ rib­ fragment,­ from­ NMMNH­ Locality­ 4507,­ in­ A,­ dorsal­ and­ B,­ posterior­ views.­ C-E, H-I,­ NMMNH­ P-61669,­Sphenacodon­sp.,­C-E,­one­complete­and­one­incomplete­dorsal­vertebra­in­C,­left­lateral,­D,­right­lateral­and­E,­anterior­views,­and­H-I,­ neural­spine,­in­H,­left­lateral and­I,­right­lateral­views,­from­NMMNH­Locality­7060.­F-G,­NMMNH­P-32233,­Sphenacodontidae,­one­neural­spine­ fragment,­from­NMMNH­Locality­4507,­in­F,­right­lateral­and­G,­left­lateral­views.­J-M,­NMMNH­P-61668,­Sphenacodontidae­indet.,­left­tibia,­from­ NMMNH­Locality­7057,­in­J,­lateral,­K, medial,­L,­proximal­and­M,­distal­views.­N-P,­NMMNH­P-61607,­Diadectidae­indet.,­incomplete­vertebra,­ from­NMMNH­Locality­4035,­in­N,­anterior,­O,­posterior­and­P,­right­lateral­views.­Scale­bars­=­A-G,­N-P, upper­left­and­H-M,­lower­right. 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­337 vertebrae,­ associated­ ribs­ and­ unidentified­ fragments­ NMMNH­ Locality­ 5381:­ NMMNH­ P-61642,­ 7­ cranial­ (Langston­1953a:­fig.­1).­From­UCMP­V3740,­the­Johnson­ fragments;­NMMNH­ P-61643,­ sacral­vertebra­fragment;­ locality:­UCMP­42158,­caudal­intercentrum. NMMNH­P-61644,­4­zygapophyses;­NMMNH­P-61645,­ two­metapodial­fragments.­From­NMMNH­Locality­7062:­ Discussion.—Langston­ (1953a:­ fig.­ 1)­ described­ and­ il- NMMNH­ P-61673,­ zygapophyses;­ NMMNH­ P-61674,­ lustrated­ six­ articulated­ embolomerous,­ dorsal­ vertebrae­ ungual­phalanx.­From­UCMP­V3740,­the­Johnson­local- preserved­ together­ with­ their­ displaced­ ribs­ in­ a­ matrix­ ity:­NMMNH­P-51876,­jaw­fragment­in­matrix­block­(Fig.­ block­of­red-mottled­micaceous­arkose­collected­from­the­ 6L),­UCMP­195511,­right­mandible­(Fig.­7L).­ Spanish­Queen­Mine­vertebrate­locality.­Although­Langs- ton­believed­UCMP­39179,­as­well­as­a­small,­embolomer- Discussion.—Second­ only­ to­ sphenacodontid­ specimens­ ous­caudal­intercentrum­from­the­Johnson­locality­(UCMP­ in­numerical­abundance,­fossils­diagnostic­of­the­herbivo- V3740),­ to­ be­ generically­ indeterminate,­ he­ suggested­ rous­family­Diadectidae­are­present­at­seven­fossil­verte- their­close­relationship­to­Archeria.­ brate­localities­near­Jemez­Springs,­and­are­present­in­all­ three­vertebrate­fossil­assemblages­of­the­local­Abo­For- mation­section.­These­specimens­included­isolated,­associ- Order­Nectridea­Miall,­1875 ated,­and­articulated­postcrania­and­demonstrate­a­general­ Family­Diplocaulidae­Cope,­1881 pattern­of­size­increase­stratigraphically,­with­the­smallest­ Diplocaulus Cope,­1877 specimens­from­assemblage­A­and­the­largest­from­assem- Diplocaulus sp. blage­C.­Further­preparation­and­study­of­this­material­will­ (Figs.­6M–N) allow­us­to­determine­with­confidence­the­number­of­di- adectid­taxa­present­in­these­assemblages. Referred specimens.—From­CM­1000:­CM­38028,­verte- bra.­From­UCMP­V3740,­the­Johnson­locality:­NMMNH­ Order­Synapsida­Osborn,­1903 P-51883,­P-61686,­vertebrae­(Figs.­6M–N). Suborder­Eupelycosauria­Kemp,­1982 Discussion.—Until­recently,­the­only­specimen­diagnostic­ Family­Ophiacodontidae­Nopsca,­1923 of­ the­ nectridean­ genus­ Diplocaulus­ recovered­ from­ the­ Ophiacodon­Marsh,­1878 Abo­Formation­near­Jemez­Springs­was­a­single­vertebra­ Ophiacodon sp. (CM­38028)­reported,­but­not­illustrated,­by­Berman­and­ (Figs.­7C–H) Reisz­(1980)­and­Berman­(1993).­This­specimen­was­col- lected­from­CM­Locality­1000­in­stratigraphic­interval­C­ Referred specimen.—From­UCMP­Locality­V3740,­the­ of­the­local­Abo­Formation­section.­Recent­processing­of­ Johnson­locality:­UCMP­195505,­195506,­195507,­dorsal­ heavily­indurated­limestone­matrix­from­the­stratigraphical- vertebrae­(Figs.­7C–H). ly­lowest­fossil­vertebrate­locality­in­Cañon­de­San­Diego­ (UCMP­V3740,­the­Johnson­locality)­has­produced­two­ad- Discussion.—The­Johnson­locality­is­the­only­site­in­the­ ditional­Diplocaulus­vertebrae­(Figs.­6M–N).­These­small,­ Abo­ Formation­ near­ Jemez­ Springs­ from­ which­ speci- holospondylous­vertebrae­are­characterized­by­an­elongate­ mens­ of­ the­ eupelycosaur­ genus­ Ophiacodon­ have­ been­ spool-shaped­centrum,­and­a­low,­rounded­neural­spine,­fea- recovered.­This­material­consists­of­small­vertebrae­with­ tures­diagnostic­of­the­nectridean­genus­Diplocaulus­(Case­ characteristic­wing-like­transverse­processes­(Figs.­7C–H)­ 1911;­Mehl­1926;­Milner­1996;­Harris­et­al.­2005). that­were­collected,­but­not­reported,­by­a­party­from­the­ University­of­California­in­1937­(Langston­1953b).­ Order­Diadectomorpha­Watson,­1917 ­Family­Sphenacodontidae­Williston,­1912 Family­Diadectidae­Cope,­1880 Genus­Sphenacodon Marsh,­1878 Diadectidae indet.­genus­and­species Sphenacodon ferox Marsh,­1878 (Figs.­6L,­7L,­9N–P) (Figs.­10–11) Referred specimens.—From­ CM­ Locality­ 1000:­ CM­ Referred specimens.—From­ NMMNH­ Locality­ 4507:­ 86173,­ vertebra.­ From­ CM­ Locality­ 1001:­ CM­ 38041,­ NMMNH­P-61689,­neural­spine­segment.­From­NMMNH­ postcranial­skeleton.­From­CM­Locality­4050:­CM­86167,­ Locality­4035:­NMMNH­P-­27474,­partial­skull;­NMMNH­ concreted­ postcrania.­ From­ NMMNH­ Locality­ 4035:­ P-61610,­ 11­ associated­ vertebrae­ and­ six­ neural­ spines;­ NMMNH­ P-61607,­ 12­ associated­ vertebrae­ and­ scrap­ NMMNH­P-61611,­numerous­associated­bone­fragments.­ (Figs.­ 9N–P);­ NMMNH­ P-61608,­ scapular­ blade.­ From­ From­NMMNH­Locality­5366:­NMMNH­P-61630,­neural­ NMMNH­Locality­4507:­NMMNH­P-32241,­ungual­pha- spine­segment;­NMMNH­P-61631,­eight­neural­spine­frag- lanx;­ NMMNH­ P-32235,­ two­ zygapophyses;­ NMMNH­ ments.­From­NMMNH­Locality­5381:­NMMNH­P-61660,­ P-32234,­centrum­fragment;­NMMNH­P-32243,­jaw­frag- P-61661,­P-61662,­P-61663,­P-61664,­P-61665,­neural­spine­ ment;­ NMMNH­ P-61612,­ neural­ spine­ fragment.­ From­ segments;­NMMNH­P-61666,­eight­neural­spine­fragments.­ 338 annaLS oF carneGie muSeum voL. 80

Fig.­10.—Sphenacodon ferox,­NMMNH­P-27474,­incomplete­skull,­from­NMMNH­Locality­4035,­in­A,­dorsal,­B,­palatal,­C,­right­lateral,­D, anterior­ and­E,­posterior­views.­F,­closeup­of­braincase.­G,­closeup­of­left­palatal­denticle­field.­Scale­bars:­A-E,­upper­right;­F,­middle­right;­and­G, lower­right.

From­ UCMP­ V3740,­ the­ Johnson­ locality:­ NMMNH­ sociated­vertebrae,­neural­spines,­ribs,­and­jaw­fragments. P-51879,­P-51880,­P-51881,­neural­spine­segments.­From­ NMMNH­Locality­7060:­NMMNH­P-61669,­numerous­as- Description.—NMMNH­P-27474­(Figs.­10–11)­is­an­in- 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­339

Fig.­11.—Interpretive­drawing­of­Sphenacodon ferox­skull,­NMMNH­P-27474,­from­NMMNH­Locality­4035,­in­A,­dorsal,­B,­palatal,­C,­right­lateral,­ and­D, anterior­views.­Broken­bone­surfaces­are­cross-hatched,­matrix­is­gray,­reconstructed­areas­are­stippled,­and­bone­impressions­in­matrix­are­ outlined­by­dashes.­Abbreviations: bo,­basioccipital;­bps,­basiparasphenoid;­de,­denticles;­ji,­jugal­impression;­mx,­maxilla;­pt,­pterygoid;­q,­quadrate,­ so,­supraocciptal;­t,­teeth;­ti,­tooth­impression; un,­unknown­bone­fragment. complete­Sphenacodon­skull­encased­in­matrix,­with­some­ but­based­on­the­right­ptergyoid­the­denticle­field­does­not­ cranial­elements­being­represented­solely­as­impressions­in­ extend­to­the­element’s­lateral­margin.­The­quadrate­pro- the­matrix.­The­right­maxilla­is­preserved­as­a­small­patch­ cesses­are­complete­and­fully­visible­in­ventral­view,­with­ of­externally­exfoliated­bone­and­some­matrix­impressions­ a­slight­color­variation­marking­their­sutural­contact­with­ in­lateral­view­on­the­block’s­right­side.­Four­teeth­are­pre- the­quadrates.­Both­quadrates­are­nearly­complete­and­well­ served­in­articulation­with­the­right­maxilla­in­apical-basal­ exposed­in­both­medial­and­lateral­views. cross-section.­ The­ maxilla-jugal­ suture­ cannot­ be­ deter- ­ The­braincase­is­incomplete;­however,­most­of­the­ven- mined,­as­the­posterior­maxilla­and­entire­jugal­are­repre- tral­elements­can­be­discerned.­Fragments­of­the­supraoc- sented­as­matrix­impressions.­ cipital­ may­ be­ preserved­ as­ linear­ outlines­ lateral­ to­ the­ ­ Both­ pterygoids­ are­ the­ most­ completely­ represented­ basioccipital­ in­ occipital­ view.­The­ dorsal­ margin­ of­ the­ elements­of­the­skull.­The­palatal­processes­meet­near­the­ basiocciptal­has­been­damaged,­while­much­of­the­ventral­ midline­and­are­visible­in­both­dorsal­and­anterior­views.­ aspect­of­this­element­is­intact.­The­sutural­surface­with­the­ The­left­transverse­flange­is­damaged,­and­no­teeth­are­pre- basipterygoid­is­clearly­discernible.­The­flange­of­the­op- served,­whereas­the­right­transverse­flange­is­incomplete,­ isthotic­contacting­the­parabasisphenoid­is­incomplete,­but­ but­preserves­at­least­four­teeth.­The­denticle­fields­on­both­ visible,­with­both­of­the­fenestrae­ovalis­readily­apparent.­ pterygoids­are­difficult­to­discern­given­the­encasing­matrix,­ The­parabasisphenoid­is­complete,­but­its­sutural­contact­ 340 annaLS oF carneGie muSeum voL. 80 with­the­quadrates­is­partially­obscured­by­damage­to­the­ disarticulated­ skeleton.­ From­ NMMNH­ Locality­ 4507:­ specimen.­ The­ basal­ tubera­ form­ triangular­ ridges­ that­ NMMNH­P-32233,­large­neural­spine;­NMMNH­P-32229,­ project­posteriorly,­and­only­the­left­carotid­foramen­is­vis- large­proximal­humerus. ible­in­this­specimen.­ ­ An­unidentified­triangular­element­is­positioned­trans- Discussion.—Romer­(1937)­and­Romer­and­Price­(1940)­ versely­in­front­of­the­temporal­fenesta­and­is­visible­in­ described­ a­ large­ species­ of­ the­ genus­ Sphenacodon,­ S. dorsal­view.­This­element­likely­represents­a­fragment­of­ ferocior,­ based­ on­ a­ well-preserved­ skull­ and­ anterior­ the­anterior­palate. vertebrae­from­the­Spanish­Queen­Mine­vertebrate­fossil­ ­ NMMNH­ P-27474­ can­ be­ assigned­ to­ Sphenacodon locality.­As­stated­above­we­believe­this­site­to­be­within­ based­on­the­collection­of­Sphenacodon­neural­spines­from­ stratigraphic­interval­C,­from­which­the­partial­skull­of­S. the­same­locality­(NMMNH­Locality­4035).­Further,­the­ ferox­(NMMNH­P-27474)­was­also­recovered.­Although­ small­overall­size­and­pterygoid­denticle­fields­that­do­not­ Romer­and­Price­(1940)­distinguished­S. ferocior­from­S. extend­to­the­element’s­lateral­margin­identify­this­skull­as­ ferox­based­mostly­on­size,­a­recent­redescription­of­the­ S. ferox,­following­Spielmann­et­al.­(2010). skull­of­S. ferox­demonstrated­a­total­of­11­cranial­charac- ters­by­which­these­two­species­can­also­be­distinguished­ Discussion.—The­ Jemez­ Springs­ fauna­ includes­ three­ (Spielmann­et­al.­2010). sphenacodontids,­S. ferox,­Sphenacodon ferocior­Romer,­ ­ Both­ species­ of­ Sphenacodon­ are­ also­ present­ in­ the­ 1940,­ and­ Dimetrodon occidentalis­ Berman,­ 1977.­ This­ faunal­ assemblage­ of­ the­ classic­ collecting­ locality­ of­ represents­ the­ only­ known­ co-occurrence­ of­ these­ three­ Arroyo­ del­Agua.­ However,­ both­ Langston­ (1953b)­ and­ eupelycosaur­ taxa.­ Sphenacodon is­ characteristic­ of­ and­ Romer­ (1960)­ argued­ that­ a­ biostratigraphic­ distinction­ restricted­ to­ the­ Coyotean­ land-vertebrate­ faunachron­ can­be­made­between­the ­main­vertebrate-fossil-producing­ (lvf)­(e.g.,­Lucas­2006),­and­is­the­common­eupelycosaur­ horizon­at­Arroyo­del­Agua­that­yields­S. ferox,­and­that­of­ in­the­Lower­Permian­of­New­Mexico,­while­Dimetrodon the­stratigraphically­higher­Anderson­quarry­from­which­ records­are­rare­throughout­the­state­(Cantrell­et­al.­2011).­ numerous­cranial­elements­of­S. ferocior­were­recovered.­ Neural­ spines­ characteristic­ of­ the­ medium-size­ sphena- Lucas­et­al.­(2005b)­thus­noted­that­the­potential­exists­to­ codontid,­S. ferox,­are­common­throughout ­the­fossil­ver- subdivide­the­tetrapod­localities­at­Arroyo­del­Agua­into­ tebrate­sites­of­Cañon­de­San­Diego.­These­records,­though­ two­biostratigraphic­assemblages­distinguished­primarily­ mostly­fragmentary,­demonstrate­an­abundance­of­S.­ferox­ by­different­species­of­Sphenacodon,­S. ferox­(lower­quar- in­the­Jemez­Springs­vertebrate­fossil­assemblages­com- ries)­and­S. ferocior­(two­upper­quarries­known­as­Ander- parable­to­that­of­the­characteristic­Coyotean­assemblage­ son­and­Cardillo).­Langston­(1953b)­and­Romer­(1960)­es- from­ the­ upper­ part­ of­ the­ El­ Cobre­ Canyon­ Formation­ timated­a­stratigraphic­interval­of­235­feet­(71.63­meters)­ (Cutler­Group)­in­the­nearby­Arroyo­del­Agua­area­(Ber- separate­occurrences­of­the­two­Sphenacodon­species­in­ man­1993;­Lucas­et­al.­2005b).­Laterally­compressed­neu- the­Arroyo­del­Agua­area.­Thus,­Romer­(1960)­suggested­ ral­spines­of­small­to­moderate­size­that­are­diagnostic­of­S. that­ a­ demonstrable­ trend­ in­ eupelycosaur­ evolution­ to- ferox­were­recovered­from­all­three­stratigraphic­intervals­ ward­increase­in­size­of­stratigraphically­younger­species­ of­the­Jemez­Abo­Formation­section.­These­occurrences,­ of­a­particular­genus­is­applicable­to­the­size­difference­be- and­ the­ absence­ of­ Seymouria­ Broili,­ 1904,­ indicate­ the­ tween­S. ferox­and­S. ferocior.­However,­the­co-occurrence­ entire­Jemez­Springs­vertebrate­assemblage­is­of­Coyotean­ of­S. ferox­and­S. ferocior­in­the­same­stratigraphic­interval­ age­(see­below­and­Lucas­2006). of­the­Abo­Formation­in­the­Jemez­Springs­area­contra- ­ In­addition­to­isolated­neural­spines,­S. ferox­is­recog- dicts­Romer’s­model­of­the­species-level­evolution­of­the­ nized­from­an­incomplete­skull­(NMMNH­P-27474)­(Figs.­ genus­Sphenacodon.­Clearly,­S. ferox­and­S. ferocior­are­ 10–11)­ and­ a­ group­ of­ associated­ vertebrae­ and­ neural­ coeval­ at­ some­ localities,­ not­ just­ temporally-successive­ spines­(NMMNH­P-61610)­from­NMMNH­Locality­4035­ species. in­assemblage­C.­Together­these­specimens­possibly­repre- sent­a­single­individual.­The­skull,­which­consists­primar- ily­of­bones­of­the­palate,­is­assigned­to­S. ferox­based­on­ Dimetrodon­Cope,­1878 its­associated­neural­spines,­relatively­small­size­and­the­ morphology­of­the­palatal­ramus­of­the­pterygoid­on­which­ Dimetrodon occidentalis Berman,­1977 the­dental­field ­does­not­extend­to­its­lateral­margin­(see­ (Figs.­8G–O) above).­­ Referred specimens.—From­ CM­ Locality­ 1000:­ CM­ 26565,­type,­partial­skeleton ­(Berman­1977,­text-figs.­1-3).­ Sphenacodon ferocior Romer,­1940 From­ CM­ Locality­ 4047:­ CM­ 86169,­ neural­ spine­ frag- ment.­From­NMMNH­Locality­5366:­NMMNH­P-61632,­ Referred specimens.—From­ Spanish­ Queen­ Mine­ ver- 61633,­ 61634,­ neural­ spine­ segments­ (Figs.­ 8G–O);­ tebrate­locality:­MCZ­1489,­holotype,­skull­and­anterior­ NMMNH­P-61635,­20­neural­spine­fragments. vertebrae­ (Romer­ and­ Price­ 1940:­ figs.­ 4F,­ 23,­ 26,­ 36,­ 59);­WM­11­(now­at­FMNH),­fragmentary­remains­of­a­ Discussion.—The­ first­ incontrovertible­ evidence­ that­ the­ 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­341

Fig.­12.—Pie­diagram­showing­abundance­of­various­vertebrate­taxa,­based­on­abundance­of­identifiable­vertebrate­fossils­(see­referred­specimens),­ from­the­Abo­Formation­of­the­Jemez­Springs­area.­N=205. genus­Dimetrodon­is­present­in­the­Abo­Formation­of­north- was­collected­(Romer­and­Price­1940;­Langston­1953b). ern­New­Mexico,­and­not­restricted­to­Lower­Permian­beds­ ­ Numerous­isolated­neural­spine­segments­of­Dimetrodon­ of­north-central­Texas­and­Oklahoma,­was­provided­by­Ber- (Figs.­8G–O)­with­the­characteristic­figure-eight­cross­sec- man­(1977).­Description­of­D.­occidentalis­was­based­on­a­ tion­are­present­at­a­second­site­(NMMNH­L-5366)­that­is­ partial­articulated­sail­and­associated­jaw­elements­collected­ also­within­assemblage­C.­However,­material­diagnostic­of­ from­a­site­high­in­the­Abo­Formation­(assemblage­C)­of­the­ Dimetrodon­is­absent­in­the­stratigraphically­lower­assem- Jemez­Springs­area­near­the­coeval­Spanish­Queen­Mine­ver- blages­A­and­B. tebrate­locality­from­which­the­type­material­of­S.­ferocior 342 annaLS oF carneGie muSeum voL. 80

Sphenacodontidae indet.­genus­and­species PALEOECOLOGY (Figs.­7A–B,­9J–M) The­vertebrate­fossil­assemblages­from­the­Abo­Formation­ Referred specimens.—From­ CM­ Locality­ 1001:­ in­the­Jemez­Springs­area­(Fig.­12)­are­broadly­similar­to­ NMMNH­ P-61637,­ jaw­ fragment;­ NMMNH­ P-61638,­ (and­correlative­to,­see­below)­most­of­the­vertebrate­fossil­ centrum;­ NMMNH­ P-61639,­ numerous­ bone­ fragments;­ assemblages­found­in­strata­of­the­El­Cobre­Canyon­Forma- NMMNH­P-61640.­From­CM­Locality­4047:­CM­86168,­ tion­of­the­Cutler­Group­in­the­Chama­basin­to­the­north,­at­ mandible.­ From­ CM­ Locality­ 4049:­ CM­ 86171,­ man- Arroyo­del­Agua­and­in­Cañon­del­Cobre­(El­Cobre­Cañon)­ dible;­CM­86172,­maxilla­and­ribs.­From­NMMNH­Lo- (Fig.­1).­Thus,­these­Coyotean-age­assemblages­contain­xe- cality­4507:­NMMNH­P-32240,­jaw­fragment;­NMMNH­ nacanth­sharks,­lungfishes,­temnospondyl­and­lepospondyl­ P-61688,­maxilla­fragment­with­teeth.­From­NMMNH­Lo- amphibians,­ anthracosaurs,­ diadectids,­ and­ (most­ diverse­ cality­5366:­NMMNH­P-61614,­atlas­centrum;­NMMNH­ and­abundant)­eupelycosaurs.­However,­the­Jemez­Springs­ P-61615,­atlas­centrum;­NMMNH­P-61616,­three­associ- assemblages,­as­currently­known,­are­less­diverse­than­the­ ated­cranial­fragments;­NMMNH­P-61617,­jaw­fragment;­ Arroyo­del­Agua­assemblage. NMMNH­ P-61618,­ tooth­ in­ jaw­ fragment;­ NMMNH­ ­ The­vertebrate­taxa­of­the­Jemez­Springs­vertebrate­as- P-61619,­tooth;­NMMNH­P-61620,­P-61621,­tooth­in­ma- semblages­represent­a­mixed­terrestrial-aquatic­fauna.­Key­ trix­block;­NMMNH­P-61622,­cervical­centrum;­NMMNH­ distinguishing­features­of­this­fauna­include­a­diverse­sphe- P-61623,­ lumbar­ centrum;­ NMMNH­ P-61624,­ P-61625,­ nacodontid­component­in­which­three­different­species­co- sacral­ vertebra;­ NMMNH­ P-61626,­ P-61627­ rib­ frag- occur­ in­ the­ same­ stratigraphic­ interval,­ and­ the­ relative­ ment;­NMMNH­P-61628,­distal­tibia;­NMMNH­P-61629,­ abundance­of­diadectid­material­combined­with­sparse­re- proximal­femur­fragment.­From­NMMNH­Locality­5381:­ mains­of­semi-terrestrial­or­semi-aquatic­vertebrates­such­as­ Eryops­and­Ophiacodon.­With­the­exception­of­Trimerorha- NMMNH­P-61646,­articular­fragment;­NMMNH­P-61647,­ chis,­specimens­of­obligatory­aquatic­amphibians­and­fish­ quadrate­ fragment;­ NMMNH­ P-61648,­ P-61649,­ lateral­ are­rare. flange­of­pterygoid;­NMMNH­P-61650,­dentigerous­ptery- ­ The­well-preserved,­articulated­condition­of­the­holotype­ goid­fragment;­NMMNH­P-61651,­maxilla­fragment­with­ of­ Trimerorhachis­ sandovalensis­ from­ the­ Jemez­ Springs­ two­teeth;­NMMNH­P-61652,­six­jaw­fragments;­NMMNH­ area­is­in­direct­contrast­to­the­sparse,­fragmentary­remains­ P-61653,­ dentigerous­ dentary­ fragments;­ NMMNH­ of­other­obligatory­aquatic­amphibians­of­the­Jemez­Springs­ P-61654,­three­tooth­fragments;­NMMNH­P-61655,­ver- vertebrate­fossil­assemblages­such­as­Diplocaulus,­Archeria,­ tebra;­ NMMNH­ P-61656,­ crushed­ vertebra;­ NMMNH­ and­Zatrachys.­The­poor­representation­of­this­component­ P-61657,­ 10­ vertebral­ fragments;­ NMMNH­ P-61658,­ of­the­fauna,­coupled­with­the­absence­of­rhipidistian­fish,­ pelvic­ girdle­ fragment;­ NMMNH­ P-61659,­ two­ metapo- apparently­represents­a­preservational­bias­against­aquatic­ dial­fragments.­From­NMMNH­Locality­7057:­NMMNH­ taxa.­Or,­it­may­indicate­that­there­were­not­areas­of­perma- P-61668,­complete­left­tibia­(Figs.­9J-M).­From­NMMNH­ nent­water­bodies­of­sufficient­size­to­support­populations­of­ Locality­ 7062:­ NMMNH­ P-61675,­ P-61676,­ vertebra;­ these­animals. NMMNH­ P-61677,­ three­ caudal­ vertebrae;­ NMMNH­ ­ Jemez­Springs­is­one­of­five­Diplocaulus­sites­in­Lower­ P-61678,­ 11­ vertebral­ fragments;­ NMMNH­ P-61679,­ Permian­red-bed­sediments­of­mostly­stream-channel­depo- proximal­central­of­carpus;­NMMNH­P-61680,­metatarsal­ sition­in­the­Abo­and­Sangre­de­Cristo­formations­of­New­ I;­NMMNH­P-61681,­seven­metapodial­fragments.­From­ Mexico.­However,­Diplocaulus­fossils­are­absent­from­up- UCMP­V3740,­the­Johnson­locality:­NMMNH­P-51877,­ per­Virgilian­and­lower­Wolfcampian­tetrapod­assemblages­ transverse­flange­of­pterygoid;­NMMNH­P-51878,­tooth­in­ of­New­Mexico­deposited­in­mixed­marine-nonmarine­strata­ matrix;­UCMP­195504,­right­femur­(Figs.­7A–B);­unnum- in­which­fossils­of­paleoniscoid­fish­have­been­found­(Red­ bered­ UCMP­ specimens,­ incomplete­ dentary,­ quadrate,­ Tanks­ and­ Laborcita­ members­ of­ the­ Bursum­ Formation:­ proximal­and­distal­humeri,­proximal­and­distal­femora. Harris­et­al.­2004,­2005).­Thus,­the­distribution­of­Diplocau- lus­fossils­in­New­Mexico­suggests­a­habitat­preference­for­ Discussion.—The­ fossil­ vertebrate­ localities­ of­ the­Abo­ paleoenvironments­subject­to­seasonal­drought.­ Formation­in­the­Jemez­Springs­area­yield­numerous­iso- ­ The­rarity­of­fossils­of­the­eupelycosaur­Ophiacodon­in­ lated,­mostly­fragmentary­elements­with­features­character- the­ Jemez­ Springs­ vertebrate­ fossil­ assemblages­ contrasts­ istic­of­the­Sphenacodontidae­(Figs.­7A–B,­9J–M).­How- with­ their­ abundance­ in­ other­ well-known­ fossil­ tetrapod­ ever,­sphenacodontid­cranial­and­postcranial­morphology­ assemblages­ near­ the­ Pennsylvanian-Permian­ boundary­ is­remarkably­conservative­so­that­the­generic­distinction­ (e.g.,­Kissel­and­Lehman­2002;­Lucas­et­al.­2005a,­2005b,­ between­Sphenacodon­and­Dimetrodon­is­based­almost­en- 2010),­as­well­as­in­the­nearby­characteristic­Coyotean­as- tirely­on­neural­spine­morphology.­Nevertheless,­the­great­ semblage­in­the­Arroyo­del­Agua­area­of­Rio­Arriba­County­ number­of­indeterminate­sphenacodontid­elements­recov- (Lucas­et­al.­2005b;­Lucas­2006)­and­a­principal­correlative­ ered­from­nearly­every­fossil­vertebrate­site­in­the­Jemez­ in­the­Halgaito­Formation­of­the­Arizona-Utah­borderland­ Springs­ area­ demonstrates­ the­ dominance­ of­ the­ Jemez­ (Sumida­et­al.­1999;­Lucas­2006).­The­paucity­of­Ophia- Springs­vertebrate­assemblages­by­Sphenacodontidae.­ codon­and­Eryops­remains­in­Abo­Formation­sediments­of­ 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­343 the­Jemez­Springs­area,­coupled­with­the­absence­of­Eda- phosaurus­ Cope,­ 1882­ (its­ characteristic­ neural­ spines,­ if­ present,­ would­ be­ easily­ recognized­ in­ the­ field),­ suggest­ an­atypically­small­semi-aquatic­or­semi-terrestrial­tetrapod­ faunal­component­in­vertebrate­assemblages­B­and­C.­The­ abundance­of­diadectid­remains,­coupled­with­the­apparent­ absence­ of­ edaphosaurids,­ suggests­ a­ paleoflora­ in­ which­ terrestrial,­high-fiber­plants­were­dominant.­ ­ The­ plants­ from­ the­ Spanish­ Queen­ Mine­ locality­ are­ consistent­ with­ the­ general­ environmental­ interpretation­ of­a­warm­climate­with­seasonal­rainfall,­indicating­a­sub- stantial­part­of­the­year­with­evapotranspiration­exceeding­ rainfall­on­a­monthly­basis.­The­paleoflora­includes­plants­ that­ have­ relatively­ broad­ leaves­ and­ probably­ moderate­ demands­for­soil­moisture­(higher­than­cactus­or­other­arid­ land­plants).­These­Abo­plants­had­woody­petioles­and­were­ likely­adapted­to­shedding­their­leaves­seasonally,­similar­ to­ocotillo­or­other­such­modern­broad-leaved­plants­of­arid­ environments. ­ The­co-occurrence­of­the­sphenacodontid­genera­Dime- trodon­and­Sphenacodon­is­rare,­with­only­three­other­co- Fig.­ 13.—Summary­ of­ lithostratigraphy,­ biostratigraphy,­ age,­ and­ cor- occurrences­known,­based­mostly­on­isolated­vertebrae­and­ relation­of­Abo­Formation­and­adjacent­strata­in­the­Jemez­Springs­area. neural­spines­(Vaughn­1969;­Sumida­et­al.­1999;­Harris­et­al.­ 2004).­Among­species­of­the­genus­Dimetrodon,­D. occiden- between­the­red­beds­of­the­Cutler­Group­and­the­Abo­For- talis­ranks­as­one­of­the­smallest,­with­an­estimated­weight­ mation­in­northern­New­Mexico­(Eberth­and­Miall­1991;­ of­41­kg­(Berman­1977,­2003),­whereas­S.­ferocior­was­as­ Eberth­and­Berman­1993;­Krainer­and­Lucas­2010).­For­ much­as­three­times­larger,­weighing­an­estimated­129­kg­ this­reason,­it­is­impossible­to­draw­a­clearcut­stratigraph- (Romer­and­Price­1940).­Clearly,­unlike­most­large­Dime- ic­boundary­between­the­Cutler­Group­and­the­Abo­For- trodon­species­of­the­late­Wolfcampian­and­Leonardian,­D. mation­in­northern­New­Mexico.­Indeed,­in­the­northern­ occidentalis­did­not­occupy­the­ecological­role­of­dominant­ Nacimiento­Mountains,­Wood­and­Northrop­(1946)­arbi- carnivore­in­this­fauna.­Instead,­its­diminutive­size­suggests­ trarily­separated­the­Abo­from­the­Cutler­red­beds­along­the­ predation­ on­ small­ aquatic­ amphibians­ and­ fish,­ leaving­ border­between­Rio­Arriba­and­Sandoval­counties.­Baars­ semi-aquatic­ vertebrates­ of­ moderate­ size,­ in­ addition­ to­ (1983)­stated­that­the­arkosic­red­beds­of­the­Abo­Forma- individuals­of­its­own­species,­as­potential­prey­of­S. fero- tion­are­similar­in­physical­aspect­and­age­to­the­lower­Cut- cior.­However,­with­an­estimated­weight­of­52­kg­(Romer­ ler­beds­of­the­Defiance­uplift­and­Monument­Valley­area­ 1960),­S. ferox­was­only­slightly­larger­than­D. occidentalis.­ of­the­Four­Corners­region.­We­therefore­include­the­Abo­ Therefore,­in­light­of­their­virtually­indistinguishable­skel- Formation­in­the­Cutler­Group,­as­did­Krainer­and­Lucas­ etal­morphology­(with­the­exception­of­the­dorsal­sail),­it­is­ (2010). reasonable­to­conclude­that­D. occidentalis­and­S. ferox­were­ ­ At­ many­ places­ in­ central­ New­ Mexico,­ south­­ of­ the highly­competitive­ecological­equivalents­(vicars). Jemez­Springs­area,­red­beds­of­the­Abo­Formation­discon- formably­overlie­the­Bursum­Formation­(e.g.,­Lucas­and­ BIOSTRATIGRAPHY­AND­AGE­ Krainer­2004;­Krainer­and­Lucas­2009).­Locally,­the­Abo­ Formation­base­is­a­conglomerate­that­erosively­overlies­ Four­lines­of­evidence­can­be­brought­to­bear­on­the­age­of­ the­Bursum­Formation.­Limestone­clasts­in­this­conglom- the­Abo­Formation­in­the­Jemez­Springs­area:­(1)­regional­ erate­indicate­that­substantial­parts­of­the­Bursum­Forma- lithostratigraphic­correlation;­(2)­the­age­of­the­Guadalupe­ tion­were­eroded­prior­to­deposition­of­the­Abo­red­beds,­ Box­Formation,­which­stratigraphically­underlies­the­Abo­ probably­ due­ to­ the­ formation­ of­ deeply­ incised­ valleys­ Formation;­ (3)­ fossil­ plant­ biostratigraphy;­ and­ (4)­ verte- (Krainer­and­Lucas­2009).­This­unconformity­is­also­rec- brate­ biostratigraphy.­ Here,­ we­ review­ these­ data­ to­ con- ognized­at­other­locations­and­indicates­a­major­tectonic­ clude­that­the­Abo­Formation­in­the­Jemez­Springs­area­is­ pulse­ near­ the­Virgilian/Wolfcampian­boundary­ (Krainer­ very­close­in­age­to­the­base­of­the­Permian­(Fig.­13). and­Lucas­2009;­Lucas­et­al.­2009). ­ There­is­much­evidence­that­the­unconformity­on­top­ Regional­Lithostratigraphy of­the­Bursum­Formation­resulted­from­a­major­tectonic­ pulse­of­the­ancestral­Rocky­Mountain­deformation,­caus- A­ comparison­ of­ the­ lithofacies­ and­ mineralogical­ com- ing­a­significant­rejuvenation­of­basement­uplifts­that­re- position­demonstrates­that­no­significant­differences­exist­ sulted­in­increased­siliciclastic­influx­and­the­deposition­of­ 344 annaLS oF carneGie muSeum voL. 80 nonmarine­red­beds­of­the­Abo­Formation­in­central­New­ Triticites­ (comparable­ with­ the­Tethyan­ genus­ or­ subge- Mexico.­However,­in­the­Chama­basin­this­unconformity­ nus­Schwageriniformis).­In­the­Jemez­Springs­section­of­ near­ the­Virgilian/Wolfcampian­boundary­ is­ not­ obvious­ Krainer­et­al.­(2005)­specimens­of­Triticites­(Triticites)­ex­ within­the­Cutler­Group­in­the­upper­part­of­the­El­Cobre­ gr.­rhodesi­Needham,­1937,­are­numerous­near­the­top­of­ Canyon­Formation­(Lucas­and­Krainer­2005a,­2005b).­In­ the­ Jemez­ Springs­ Member­ and­ indicate­ a­ late­Virgilian­ the­Jemez­Springs­area­siliciclastic­red­beds­of­nonmarine­ age­(=­late­Gzhelian;­=­VC3­biozone­of­Wilde­1990).­This­ origin­ at­ the­ base­ of­ the­Abo­ Formation­ rest­ directly­ on­ places­a­maximum­age­of­late­Virgilian­on­the­base­of­the­ marine­strata­at­the­top­of­the­Guadalupe­Box­Formation.­ Abo­Formation­in­the­Jemez­Springs­area­(Fig.­13). Our­ lithostratigraphic­ data­ (Fig.­ 4)­ suggest­ a­ substantial­ amount­of­stratigraphic­relief­(>60­m)­in­the­Jemez­Springs­ Fossil­Plant­Biostratigraphy area.­It­thus­seems­likely­that­there­is­an­unconformity­at­ the­base­of­the­Abo­Formation­in­the­Jemez­Springs­area.­ The­Spanish­Queen­Mine­paleoflora­is­typical­of­Read­and­ Available­data,­however,­do­not­allow­a­precise­estimate­ Mamay’s­ (1964)­ Floral­ Zone­ 13,­ the­ Zone­ of­ Callipter- of­the­temporal­hiatus­represented­by­the­unconformity.­It­ is­ Brongniart,­ 1849,­ which­ they­ place­ at­ the­ base­ of­ the­ probably­is­a­hiatus­between­middle­Wolfcampian­(age­of­ Permian.­ However,­ with­ the­ recently­ repositioned­ Penn- base­of­Abo­Formation­regionally)­and­late­Virgilian­strata,­ sylvanian-Permian­ boundary­ to­ within­ the­Wolfcampian,­ but­this­cannot­be­demonstrated­with­certainty. this­zone­straddles­the­Pennsylvanian-Permian­boundary.­ Stratigraphically,­the­ Spanish­ Queen­ Mine­ paleoflora­ is­ Age­of­the­Guadalupe­Box­Formation similar­to­that­of­the­Lower­Rotliegend­of­Germany­(Kerp­ and­ Fichter­ 1985;­ Barthel­ 2009),­ particularly­ the­ lower­ Henbest­and­Read­(1944),­Read­and­Wood­(1947),­DuCh- portions­ of­ that­ interval,­ where­ a­ secure­ Pennsylvanian- ene­ (1973),­ Swenson­ (1996),­ Kues­ (1996,­ 2001),­ and­ Permian­ position­ is­ uncertain.­Taxa­ of­ particular­ note­ in­ Krainer­et­al.­(2005)­are­among­those­who­have­reviewed­ making­ this­ determination­ include­ Neurodontopteris au- previous­studies­of­the­lithostratigraphy­and­biostratigra- riculata,­ Ernestiodendron filiciforme,­ Culmitzschia/Le- phy­of­the­Pennsylvanian­marine­strata­in­the­Jemez­Moun- bachia speciosa,­Pecopteris bredovii­Germar,­1845, and­ tains.­ In­ the­ Jemez­ Springs­ area,­ the­ dominantly­ marine­ Gomphostrobus bifidus­(Geinitz)­Potonié,­1891,­all­primar- Pennsylvanian­stratigraphic­interval­immediately­beneath­ ily­Late­Pennsylvanian­and­Early­Permian­in­their­occur- the­Abo­Formation­belongs­to­the­Guadalupe­Box­Forma- rences.­However,­the­Spanish­Queen­Mine­paleoflora­also­ tion,­ which­ can­ be­ divided­ into­ two­ members,­ an­ upper­ contains­ elements,­ such­ as­ conifers,­ Dichophyllum­ Elias­ Jemez­Springs­Member­(Sutherland­and­Harlow­1967)­and­ ex­Andrews,­1941,­and­Taeniopteris,­reported­together­as­ a­lower­San­Diego­Canyon­Member­(Krainer­et­al.­2005).­ early­as­the­Missourian­from­Midcontinent­North­America­ ­ The­Jemez­Springs­Member,­particularly­in­the­upper­ (Moore­ et­ al.­ 1936;­ Cridland­ and­ Morris­ 1963)­ and­ the­ part,­ contains­ abundant­ marine­ fossils,­ mostly­ well-pre- Appalachians­(McComas­1988).­Other­publications­have­ served­ brachiopods,­ gastropods,­ bivalves,­ crinoids,­ and­ shown­that­plants­such­as­conifers­may­occur­in­the­Middle­ bryozoans,­indicating­a­low-energy­shallow­marine­envi- Pennsylvanian­of­the­American­Illinois­Basin­(Plotnik­et­ ronment­ with­ fine-grained­ siliciclastic­ influx­ (Sutherland­ al.­2009;­Falcon-Lang­et­al.­2009);­conifer­occurrences­in­ and­Harlow­1967;­Kues­1996;­Swenson­1996;­Krainer­et­ the­Middle­Pennsylvanian­are­well­established­in­Europe­ al.­2005).­The­limestone­beds,­which­contain­a­diverse­in- (e.g.,­Galtier­et­al.­1992,­among­many­other­reports).­In­all­ vertebrate­fossil­assemblage,­were­deposited­during­short­ these­instances,­early­appearances­of­plants­such­as­those­ periods­of­reduced­siliciclastic­influx­in­a­shallow,­open- dominant­ at­ the­ Spanish­ Queen­ Mine­ locality,­ coincide­ marine­setting.­Muddy­textures­(mostly­wackestone)­indi- with­ paleoenvironments­ indicative­ of­ seasonally­ dry­ cli- cate­low­energy­conditions.­A­few­limestone­beds­contain­ matic­conditions­and­associated­seasonal­moisture­stress.­ age-diagnostic­fusulinids­(Krainer­et­al.­2005). ­ At­present­there­are­too­few ­occurrences­of­these­sea- ­ The­base­of­the­Guadalupe­Box­Formation­is­thus­dated­ sonally­dry­paleofloras­to­allow­the­high­precision­needed­ by­ fusulinid­ biostratigraphy­ as­ latest­ Moscovian­ (=­ late­ for­refined­biostratigraphic­interpretations.­An­example­of­ Desmoinesian,­DS4­zone­of­Wilde­1990)­because­of­the­ the­problems­that­can­result­from­this­is­the­study­of­Wag- presence­ of­ Fusulina­ aff. F. acme­ Dunbar­ et­ al.,­ 1942,­ ner­and­Lyons­(1997),­who­incorrectly­interpreted­the­7-11­ accompanied­ by­ Climacammina­ ex­ gr.­ magna­ Roth­ and­ flora­from­the­Missourian­of­Ohio­described­by­McComas­ Skinner,­ 1930,­ and­ indeterminate­ species­ of­ Palaeotex- (1988)­ to­ be­ a­ latest­ Stephanian–equivalent­ (Gzhelian)­ tularia­Schubert,­1921,­and­Eugonophyllum­Konishi­and­ rather­than­early­Stephanian-equivalent­(Kasimovian)­age.­ Wray,­ 1961­ (see­ Krainer­ et­ al.­ 2005­ for­ documentation­ This­requires­a­gap­of­many­millions­of­years­to­exist­in­ of­these­microfossils).­The­middle­part­of­the­Guadalupe­ the­North­American­geological­section,­in­contravention­to­ Box­ Formation­ contains­ Triticites­(Schwageriniformis?)­ all­other­means­of­correlation­(Falcon-Lang­et­al.­2011).­ aff.­T.­jemezensis­Needham,­1937.­This­level­corresponds­ Similarly,­the­age­of­the­non-marine­Dunkard­Group­in­the­ to­the­Missourian­(=­Kasimovian)­MC1­or­MC2­biozones­ northern­Appalachians­has­been­difficult­to­resolve­because­ of­Wilde­(1990)­due­to­the­primitive­stage­of­evolution­of­ of­the­intercalation­of­Permian-like,­callipterid­and­conifer- 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­345 rich­floras­with­Pennsylvanian-like­wetland­floras­(Darrah­ lithologic­ similarity­ and­ similar­ stratigraphic­ position­ of­ 1975).­Fortunately,­in­the­case­of­the­Spanish­Queen­Mine­ the­lower­part­of­the­Abo­Formation­in­the­Jemez­Springs­ paleoflora,­there­are­species­that­point­clearly­to­a­latest­ area­and­the­upper­part­of­the­El­Cobre­Canyon­Forma- Pennsylvanian­or­earliest­Permian­age,­such­as­Alethopter- tion­in­the­Chama­basin.­Thus,­the­lower-middle­part­of­ is schneideri­Sterzel,­1881,­Callipteridium gigas­(Gutbier)­ the­Abo­Formation­and­its­fossil­assemblages­in­the­Jemez­ Weiss­1870,­Neurodontopteris auriculata,­Rhachiphyllum Springs­area­are­best­assigned­a­middle­Wolfcampian­age,­ schenkii,­and­Culmitzschia/Lebachia speciosa,­consistent­ close­to­the­Pennsylvanian-Permian­boundary­(Fig.­13).­ with­other­lines­of­evidence­for­the­age­of­these­rocks. ACKNOWLEDGMENTS

Vertebrate­Biostratigraphy We­are­grateful­to­the­U.S.­Forest­Service­and­several­private­landowners­ for­access­to­Abo­Formation­outcrops­in­the­Jemez­Springs­area.­For­as- ­ In­the­Chama­basin,­particularly­at­Cañon­del­Cobre,­ sistance­in­the­field­we­acknowledge­J.­Hunley,­A.­Lerner,­K.­Madalena,­ the­lower­part­of­the­Cutler­Group­(El­Cobre­Canyon­For- P.­Reser,­and­several­NMMNH­volunteers.­P.­Neville­generously­donated­ mation)­ yields­ plant­ and­ vertebrate­ fossils­ of­ both­ Late­ fossils­he­found­near­Jemez­Springs­to­the­NMMNH.­P.­Holroyd­facili- Pennsylvanian­and­Early­Permian­age­(Lucas­et­al.­2010).­ tated­access­to­the­UCMP­collection­and­provided­photographs,­and­C.­ Norris­provided­information­on­and­photographs­of­the­YPM­collection.­ At­Arroyo­del­Agua­only­the­upper­part­of­the­El­Cobre­ The­comments­of­two­anonymous­reviewers­improved­the­content­and­ Canyon­ Formation­ is­ exposed,­ and­ the­ vertebrate­ fossil­ clarity­of­the­manuscript. assemblages­are­of­Early­Permian­(middle­Wolfcampian)­ age­(Lucas­2006).­Indeed,­west­of­Arroyo­del­Agua,­near­ LITERATURE­CITED Jaralosa­and­Gallina,­marine­limestones­of­the­upper­part­ of­the­Guadalupe­Box­Formation­yield­Virgilian­fusulinids­ BaarS, D.L. 1983.­The­Colorado­Plateau.­A­Geologic­History.­University­ (Henbest­and­Read­1944),­and­these­fusulinids­are­strati- of­New­Mexico­Press,­Albuquerque.­279­pp. graphically­below­the­Coyotean-age­vertebrate­fossil­as- BartHeL, m.­ 2009.­ Die­ Rotliegendflora­ des­ Thüringer­ Waldes.­ Sonderveröffentlichung­ Naturhistorisches­ Museum,­ Schleusingen.­ semblages­at­Arroyo­del­Agua.­ 165­pp.­ ­ In­the­Chama­basin,­the­lowest­occurrence­of­the­eupe- Berman, D.S­1977.­A­new­species­of­Dimetrodon­(Reptilia:­Pelycosauria)­ lycosaur­Sphenacodon­marks­the­beginning­of­the­Coyo- from­the­Lower­Permian­of­north-central­New­Mexico.­Journal­of­ tean­land-vertebrate­faunachron,­which­began­during­the­ Paleontology,­51:108–115. ———.­1993.­Lower­Permian­vertebrate­localities­of­New­Mexico­and­ latest­Pennsylvanian­and­encompasses­much­of­Wolfcam- their­ assemblages.­ New­ Mexico­ Museum­ of­ Natural­ History­ and­ pian­time­(Lucas­2006).­The­vertebrate­fossil­assemblages­ Science­Bulletin,­2:11–21. from­the­Abo­Formation­in­the­Jemez­Springs­area­clearly­ ———.­ 2003.­ New­ materials­ of­ Dimetrodon teutonis­ (Synapsida:­ are­of­Coyotean­age.­This­indicates­correlation­of­the­low- Sphenacodontidae)­from­the­Lower­Permian­of­Germany.­Annals­of­ er-middle­Abo­Formation­at­Jemez­Springs­with­the­upper­ Carnegie­Museum,­73:108–116. 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appenDix 1: Coordinates­Of­Measured­Sections Map­coordinates­of­measured­sections­(section­location:­Fig.­2;­stratigraphic­sections:­Fig.­4)­ (UTM­coordinates,­datum­NAD­27,­all­zone­13N).

Spanish Queen Mine —­base­at­345190E,­3955315N;­top­at­345611E,­3955122N.­ Neville —­base­at­344672E,­3954765N;­top­at­344622E,­3954931N. Vista Linda —­base­at­344473E,­3953530N;­top­at­344858E,­3953391N. Dimetrodon Site (= Jemez River section of Krainer and Lucas 2010) — base­at­343754E,­3952554N;­top­at­344598E,­3952583N. Mesa de los Datiles — base­at­344855E,­3953389N;­top­at­342989E,­3949806N. Gilman B — base­at­343003E,­3949723N;­top­at­343954E,­3950169N. Gilman A —­base­at­341134E,­3952490N;­top­at­341224E,­3952720N. 2012­ LucaS et aL.—upper paLeozoic aBo Formation ­­­­­­­­­­­­­­­349

appenDix 2:­Vertebrate­Taxa­By­Locality­In­The­Abo­Formation In­The­Jemez­Springs­Area­(continued on the next page.)

ASSEMBLAGE A UCMP Locality V3740 [Johnson locality] (= NMMNH L-6842) Eryops sp.­(vertebrae,­UCMP­specimens;­dermal­bone,­UCMP­195508;­partial­palate­with­teeth,­UCMP­195509;­premaxilla­with­two­large­teeth,­ UCMP­195510) Embolomeri­indet.(embolomerous­vertebra,­UCMP­42158)­(see­Langston­1953a,­1953b) Diplocaulus­sp.­(vertebra,­NMMNH­P-51883;­vertebra,­NMMNH­P-61686­) Diadectidae­indet. (jaw­fragment­in­matrix,­NMMNH­P-51876) Small diadectid (specimens­reported­by­Langston­1953b,­right­mandible,­UCMP­195511) Ophiacodon­sp.­(vertebrae,­UCMP­195505-195507) Sphenacodontidae,­genus­indet.­(tranverse­flange­of­pterygoid,­NMMNH­P-51877;­tooth­in­matrix,­NMMNH­P-51878;­femur,­UCMP­195504;­ proximal­and­distal­femora,­proximal­and­distal­humeri,­UCMP­specimens) Sphenacodon ferox­(neural­spine­segment,­NMMNH­P-51879;­neural­spine­segment,­NMMNH­P-51880;­2­neural­spine­fragments,­NMMNH­ P-51881)

UCMP Locality V3855 (Spanish Queen Mine vertebrate locality) ?Archeria­(articulated­vertebrae,­associated­ribs­and­unidentified­fragments,­UCMP­39179) large diadectids (see­Langston­1953b) Eryops grandis­(small,­fragmentary­skull,­UCMP­39179) Sphenacodontidae indet.­(see­Langston,­1953b) Sphenacodon ferocior (skull­ and­ anterior­ vertebrae,­ holotype,­ MCZ­ 1489;­ fragmentary­ remains­ of­ a­ disarticulated­ skeleton,­ WM­11)

ASSEMBLAGE B NMMNH L-4507 Eryops sp.­(intercentrum,­NMMNH­P-32230) Diadectidae indet.­(partial­centrum,­NMMNH­P-­32234;­zygapophyses,­NMMNH­P-­32235;­neural­spine,­NMMNH­P-61612;­ungual­phalanx,­ NMMNH­P-32241;­jaw­fragment,­NMMNH­P-32243)­ Spenacodontidae, genus indet.­(vertebra­with­ribs,­NMMNH­P-32236;­dentigerous­maxilla­fragment;­dentigerous­jaw­element,­NMMNH­P-32240;­ proximal­femur,­NMMNH­P-61687) Sphenacodon ferox­(neural­spine,­NMMNH­P-­61689) Sphenacodon ferocior (neural­spine,­NMMNH­P-­32233;­proximal­humerus,­NMMNH­P-­32229)

NMMNH L-5381 ?Eryops­sp.­(intercentrum,­NMMNH­P-61641) Diadectidae indet.­(7­cranial­fragments,­NMMNH­P­61642;­sacral­vertebra­fragment,­NMMNH­P-61643;­4­zygapophyses,­NMMNH­P-­61644;­2­ metapodial­fragments,­NMMNH­P-61645) Sphenacodon ferox (vertebra,­NMMNH­P-61660;­neural­spine­segments,­NMMNH­P-­61661­through­P-61665;­8­neural­spine­fragments,­NMMNH­ P-61666)

ASSEMBLAGE C CM Locality 999 Xenacanthus­(tooth,­CM­38026)­

CM Locality 1000 (= NMMNH L-7056) ?Eryops sp.­(vertebral­intercentrum­and­neural­spine,­CM­26567;­jaw­fragments,­CM­86177)­ Diplocaulus­sp.­(vertebra,­CM­38028),­ Zatrachys serratus­(tabular,­CM­41706),­ Diadectidae indet.­(vertebra,­CM­86173) Sphenacodon­sp.­(occiput,­CM­26566) Dimetrodon occidentalis (partial­skeleton,­holotype,­CM­26565)

CM Locality 1001 (= NMMNH L-5380) ?Eryops sp.­(skull­fragments,­CM­86175) ?Trimerorhachis sp.­(partial­skull­w/­element­of­postcrania,­CM­38027)­ Diadectidae­(most­of­skeleton­minus­skull,­CM­38041)­ Spenacodontidae, genus indet. (jaw­fragment,­NMMNH­P-61637;­centrum,­NMMNH­P-61638;­left­premaxilla,­CM­86176) Dimetrodon occidentalis (2­neural­spine­frags,­NMMNH­P-61640;­vertebral­and­neural­spine­fragments,­CM­86174)

CM Locality 1002 (= NMMNH L-7055) Trimerorhachis sandovalensis (skull­and­partial,­articulated­skeleton,­holotype,­CM­38025)

CM Locality 1003 Platyhystrix­sp.­(neural­spines,­CM­38029)­ 350 annaLS oF carneGie muSeum voL. 80

appenDix 2:­Vertebrate­Taxa­By­Locality­In­The­Abo­Formation In­The­Jemez­Springs­Area­(continued from previous page.) ASSEMBLAGE C, continued

CM Locality 4047 Sphenacodontid indet.­(mandible,­CM­86168) Dimetrodon occidentalis­(neural­spine­fragment,­CM­86169)

CM Locality 4048 Eryops sp.­(neural­spine,­CM­86170)

CM Locality 4049 Sphenacodontid indet.­(mandible,­CM­86171;­maxilla­and­ribs,­CM­86172­

CM Locality 4050 Diadectidae indet.­(partial­postcranial­skeleton,­CM­86167)

NMMNH L-4035 Diadectidae indet.­(12­associated­vertebrae,­NMMNH­P-­61607;­scapular­blade,­NMMNH­P-­61608) Sphenacodon ferox­(partial­skull,­NMMNH­P-27474;­11­associated­vertebrae­and­6­neural­spines,­NMMNH­P-61610;­associated­bone­fragments,­ NMMNH­P-61611)

NMMNH L-4508 Vertebrata indet.­(one­indeterminate­bone­fragment,­not­collected)

NMMNH L-5366 Temnospondyli­indet.­(jaw­frag,­NMMNH­P-­61613) Sphenacodontidae, genus indet.­(atlas­centrum,­NMMNH­P-61614;­atlas­intercentrum,­P-61615;­3­associated­cranial­fragments,­P-61616;­jaw­ fragment,­P-61617;­jaw­fragment­with­tooth,­P-61618;­tooth,­P-61619;­tooth­in­matrix­block,­P-61620;­cervical­centrum,­P-61622;­lumbar­centrum,­ P-61623;­sacral­vertebra,­61624;­sacral­vertebra,­P-61625;­rib­fragment,­P-61626;­rib­fragment,­P-61627;­distal­tibia,­P-61628;­proximal­femur,­ P-61629) Sphenacodon ferox (neural­spine­segment,­NMMNH­P-61630;­8­neural­spine­fragments,­NMMNH­P-61631) Dimetrodon occidentalis. (neural­spine­segment,­NMMNH­P-­61632;­neural­spine­segment,­NMMNH­P-61633;­neural­spine­segment,­NMMNH­ P-61634;­20­unassociated­neural­spine­frags,­NMMNH­P-61635)

NMMNH L-7057 Sphenacodontidae,­genus­indet.­(complete­tibia,­NMMNH­P-61668)

NMMNH L-7060 Sphenacodon ferox­(assoc.­vertebrae,­neural­spines­and­jaw­frags­of­one­individual,­NMMNH­P-61669)

NMMNH L-7061 Tetrapoda indet.­(vertebra­frag,­NMMNH­P-61670;­bone­frag,­NMMNH­P-61671)

NMMNH­L-7062 ?Eryops sp. (pleurocentrum,­NMMNH­P-61672) Diadectidae indet.­(zygapophyses,­NMMNH­P-61673;­ungula­phalanx,­NMMNH­P-61674)

YPM lungfish locality Gnathorhiza (two­tooth­plates,­YPM­8636-8637)