THE RELATIONSHIPS OF THE AMERICAN BLACK-FRUITED HAWTHORNS CRATAEGUS ERYTHROPODA, C. RIVULARIS, C. SALIGNA AND C. BRACHYACANTHA TO C. SER. DOUGLASIANAE (ROSACEAE)

14 Pages·1999·5.9 MB·English

by  J B Phipps

THE RELATIONSHIPS OFTHEAMERICAN
BLACK-FRUITEDHAWTHORNS
CRATAEGUSERYTHROPODA, C. RIVULARIS,
C. SALIGNA AND C. BRACHYACANTHA
TOC. SER. DOUGEASIANAE (ROSACEAE)
J.B.PHIPPS
Depart)}ieiitafPlantSciences
TheUniversityofWesternOntairo
London. Ontario,NCiA3B7.CANADA
ABS'lRACr
At^TOiipofblack-truitcdCn/tcU\i(//.\(Rosaccdf)fromthewesternUnitedStatesisanalysed.
Crataeguserythopocla,aspecieswithpreviouslypoorlyunderstoodaffinities,isshowntobe
closelyrelatedtoC.nviilaris.TheaffinityofC,rnmlarisandC.salignaisreconfirmed.The
threespeciesformacladesubtendedbyC.hnicljyaanithaanddifferentfromthecladecon-
tainingC'.clouglc/siiAndC.okeiiuoniiwhenanalyi^etlbyPAIJPusing38morphologicalchar-
acteristics.CrataegusrivulansandC.s<ilig)niareletrot\'pified.
Klis11MEN
SeestLidiaungrupodeCnildtg/is(Rosaceae)confrutosnegrosdeloestedelosBstados
Unidos.Crataeguserytljujjiula,unaespecieconalinidadespobrementeconocidas,semuestra
queestamuyrelacionatioconC'.ririildris.Sercconfirmac]ueC.rnaldr'isesrambienmuy
afi'naC.saligna.LastrescspeciesformanundadoencuyabaseseencuentraC.hracbyacaiitloa
yqueesdiferentedelcladoquecontienea(..doiiglasiiyC.okeiniouiicuandoseanaliza
medianteelPAUPusando38caracterfsticasmorfologicas.SelectotipificanCrataegusviviilans
yC.saligna.
Biack-fruiteclCrataegusareinaSLibstancialminorityworldwide,asalso
inNorthAmericawhereperhapssevenspeciesareblack-fruited.Numer-
ousauthorshavepointed out thesignificanceoffruitcolorin relation to
frugivordispersalandthistopichasevenreceivedalittleattentionforNorth
Americanhawthorns(e.g.Sallabanks I99.^).ItisnotknownwhetherNorth
Americanblack-fruited hawthorns formanaturalcladeandwithout un-
dertakinganexhaustiveexamination thatpotentiallyincludesmanyred-
fruitedspecies,thisfactcannotbedetermined. However,ithadappeared
tomethattheyfellintoatleasttwogroups,thefirstbeingallmembersof
seriesDoiiglasianaeiRthd.exSarg.)Rehd.(thisincludesC.doiigUisiiLindl.,
C.snksdorfii(Sarg.)KruschkeandC.okennottii].Y^.Phipps)togetherwitha
secondgroup(C.brachyacaiithaSarg.&Engelm.,C.erythropodaGreene,C.
nviihirisNutt.exTorn&A.GrayandC.salignaGreene)whichmighthave
Siix\18(3):617-660.1999
,
648 SiDA 1S(3)
somelooseinterrelationship.Itestthishypothesisoftwoclacleswithacla-
disticanalysisusing38morphologicalcharacteristics.
SeriesDouglashtnaehasbeenthesubjectofasubstantialamountofre-
cent researchand ismorenortherly than thegroupofspeciesonwhich I
focusinthispaper,withasouthernlimitofabout43°NexceptinCalifor-
niawhereitisfarthersouth.LoveandFeigen(1978)showedthatC.doiighisii
couldhybridizewiththedistantlyrelatedandintroducedC.nmnogynay^xcv^.
BrunsfeldandJohnston(1990)providedsolidgroundsforraisingC.siiksdorfii
(Sarg.)KruschkefromavarietyofC.doiiglasiitospecificrankwhileDickinson
etal.(1996)contrastedthebreedingsystemsinthe2()-stamenC.suksdorfii.
andthe 10-stamenC.douglasiitogoodeffect.Dickinsonandstudentsare
continuingtheirbiosystematicandmorphometricanalysesofsenDouglasianae.
Recently,Idescribedanewspecies,C.okennomi,fromthisgroup(Phipps&
O'Kennon1998).I3uetothisrecentactivityandthecontinuingresearches
oftheDickmsongrouponser.Do/zglasianae.Irestrictcoverageoftheseries
inthispapertothecladisticanalysismerelytodiscoverifser.Doiiglastanae
isaseparatecladefromtheotherspeciesconsidered.
Here,therefore,Idirectattentiontothemoresoutherly,andallopatric,
componentofthe'black-fruited'Crataegi,thegroupofC.rivularis,C.crythropoda,
C. saltgnaandC. hrachyacantha.Therelationshipsofthesespecieshasal-
waysbeenmuchmorecontroversial.CrataegussalignawasplacedwithC.
brachyacanthabyPalmer(1925),asuppositionthatIfollowedwithoutstudy
intheMaloidchecklist(Phipps,etal. 1990).Then,particularlyinfluenced
byfieldobservations,IrecognizeditsprobableaffinitytoC. nvularh,are-
lationshipinfactexplicitlynotedbyGreene(1896)whenhedescribedC.
saligna.WithregardtoC.erythropoda,asrecentlyasthisyear(Phipps1999),
I included thisspeciesamongthewestern red-fruited species.Neverthe-
less,theexistenceofafewintermediatespecimensnotedsincethatpaperwas
prepared,togetherwiththedetailedexaminationforthispaper,andprepa-
rationofthedraftdescriptionfortheFloraofNorthAmerica,showedcon-
clusivelythatitwasactuallyclosetoC.rivularis.Alsorecently,Welsh(1982)
placedbothC.rivularisandC.salignaunderC.douglasii,asolutionthat1do
notbelievethatanycurrentstudentofCrataeguswouldconsidertenable.
Nor1tthheArmeefroriecarnesbollavcekd-ftrouiitnevdesspteicgiaetseatnhde,icnlatdhiestciacseisnotfeCrreslaaltiigonan,shCi.prsivouflatrhies
andC. erythropoda,whichhaveneverreceivedanyindependenttaxonomic
studysincetheiroriginaldescriptions,incontrasttoser.Douglasianae,to
carefullycharacterizeandtypifythem.Inaddition,C.rivularisandC.saligna
areprovidedwithdetaileddistributionmapsforthefirsttime. Finally, I
needtocommentonmycautious'blackfruited'.Speciesinthisgroupeither
haveblack,purple,orburgundyfruitwhenfullyripe,theparticularcolor.
Phipps,Black-truicedCrataegusfromthewesternUnitedStates 649
anditschangesduringripening,havingcaxonomicsignificance.Theterm
'black-fruited'inthetitlethereforereferstoagroupofrelatedhawthorns
chatarepredominantly,butnotentirely,black-fruitedatfullmaturity.
Miniloos
Anextensivesampleofherbarium specimensofthe taxatobestudied
hasbeenassembledbymyownfieldworkinrecentyears,assistedbyR.J.
O'Kennon,togetherwithmanyexamplesofloanspecimens.Asthispaper
isnotaimedatassessingspecificlimits,nomorphometricpheneticanaly-
sisoflargesampleswasconducted.However,thesampleswereusedtode-
terminethecharacteristicsofsyntheticOTU's,oneforeachspecies, that
wouldbeanalyzedbyPAUP2.4.1.The38characteristicsusedforthispurpose
arelistedinTable 1.Distributionmapsforpreviouslyunmappedorinad-
cc]uatelymappedspecieswerecreated byRange-JVIapper,aprogram cre-
atedbythefirmTundraVoleofFairbanks,Alaska.Forthispurpose,where
sufficientlypreciselocationdataexisted on theherbarium labelthiswas
convertedintolatitude/longitudecoordinatesaccuratetothenearestminute.
Filesforeach specieswerethen mapped. Typification followed standard
procedures.Syntypesorpotentiallectotypeswerelocated,occasionallywith
greatdifficulty,andhololectotypesorneotypesselectedonthebasisofgoodness
offittotheprotologue,citationintheprotologue(ifpertinent),andspeci-
menquality,whenachoicewasavailable.Nospecimensareillustrateddue
totheexistenceofexcellentillustrationselsewhere.Thecharacterizations
presentedhereofthespeciesoutsidesenDouglasianaeareinfactsomewhat
detaileddiagnosesbecausethisworkdocsnotpurporttobeacaxonomic
revision,becauseexcellentdescriptionsexistelsewhereandbecause,mmy
view,theyrepresentfourwell-markedspeciesevenbyconservativecriteria.
INTHRRRLATK^NSHIPS
ThepossibleandintriguingrelationshipbetweenC.malignaandC.rivularis
hasalreadybeensuggested.Ifthesetwoprovedtobesisterspeciesthena
newseries'Rivulares couldbecreatedtoaccomodatethem. However,on
reflection,itseemedthatC rivularismightbeevenclosertoC.erythropoda
onthebasisofidenticalfloralandinflorescencecharacteristicsandfruitdiffering
onlyinfullyripecolor.AseriesthatincludedbothC.saligmiandC.erythropoda
isperhapsalittlebroaderthanmostseriesinNorthAmericanCrataegus.
WeneedalsotopayattentiontoPalmer's(1925)viewthatC.salignawas
closelyrelatedtoC.hrachyacanthaandthereforeshouldbeplacedinsenBrevispinae
(Palmerused thenomennudumBrachyacanthae),whereIhadprovisionally
placedit(Phippsetal.1990)followingconventionandwithoutstudy.Crataegus
hrachyacanthaisverysimilartoC.salignainflowerandleafcharacteristics
..
650 SiDA18(3)
Tahi.I'I.'riiirt\--L-it;lumorpholoiiKalchuriKlcristicsscaredkirclatlisticanalysis.
Plilllt.l^ii/CI\// 21. Lealteeth:glands
1 I^lantliabit
2. Barklype 22. Flower:ntimber
3- Braiuliarrant;cmcnt 23. Pedicel:pubesceiuf
'I'lmnnaiullu'ip 24. Anthesistime
4. 'I'liorns:mdeterminarcpresent.-' Flower
"S Tlioriis:leiit^ch 23. Diameter
6975..,. TTTTwllhiiioLoorirrnsnns,ss,:,1cccyioorllioorrrov::ladn,,biLr;rrccaoo\w'lonrs:browns 22229768.... CSSAtarnytlalhymeexe:rn:nl:otbcineomosl.b:oertmarums
|{). Twi^us,1yrold,color;i^'ray
L11eI\Il11871.56n1i.v....,j, LLLLIiLL.aaa.aaaaianmmmmnnmniiiiinninnnnniiiiaaaaaaaa::::::::vpvlmiklteooeo-anisbbn^nxtnieeag.^stttrsiinihsitohoui/onannbmt,prsbioeicfsnenattlrwdiiiirsemdephbstelesirt(L|ilalr)t I333333'iir'31267TOii.-.....i.i!PFIFFIFIrr'rrr'yrttttttrttiiiitieiiiiiuittttttnt:::::::essccscpl:ooahheulllaanboloyppgerarxeet,ts,,hecaormtera21(onlihhencienfletialtghcaherttsibi)peoetnnoerseskillripeness
19. Lamina:aba.xialpubescence,\'oiint; Aiitiinnhdjol'ui^c
20. Petiole:"lands 3<S.Color
butdiffersgreatlyinthethorns(ctirvec),=S1.5cmlong),coloroiovcrmattire
inflorescence(yellowish orange), fruit size(generally larger), smooth lat-
eralfacesofthepyrenes,andlackoicopper-coloredhark.AlsoWelshhad
madebothCsaliii^iu/andCrividarisvarietiesofC.cloj/iihis/i,inwhatseemed
tometobeanuntenableassociation.Itthereforeseemedpointfultocon-
ductanumericaltaxonomicstudytothrowlightonthesecontrastingpos-
sibilitiesandtoestablishwhetheranyofthesetaxawereespeciallycloseto
membersofset.Doiiglasianae.
Forthis,Iscored 38characters(Table I)for9syntheticOTUs,onefor
eachspecies,whichincludedalltheblack-fruitedCnitdci^i/ssjieciesthatare
thepartictilarsubjectsofthispaper(C..u/I/g/k/,C. rivz/laris,C.erythropoda
saenr.dDCo.ughlrdcsiiidbyUicitaen/atslhwie)llpharsstCherlIeod/-zfi^rInciitsHecsl.s.ou,tCg.rosLiiipkssdCo.rjiuiiaoUnids(CT.oronken&iiAo.niGir.o)f
ScheeleandC. )ii()iv){i_yuci}-ac<.\.
UsingPAUP2.-1.I,(].crythrojiodciandC. r'lViiUir'isareshowntobesister
speciesinthethreeshortesttrees(Fig. 1).Thedoii{\Ltsiigronp,consisting
ofCr.<:/w//i;/;/.v//andokeunini'n.alwaysformedonestib-cladewhileC.hracbyacantha,
CScdii^ihi,C.rii'/zlarisandC.erydirnpadcialwaysformedanotherinallshortest
cladograms. However, C. Scd/i^iu/and C, hrachyacanthciwere by no means
veryclosemthesecladogramsalthoughC.nviihirisandC.i^ryllintpodawere
Puipps,Black-frLiicedCrataegusfromthewesternUnitedStates 65
monogyna PAIJP2.4.1
douglasii 9OTUs
38characters
okennonii
' suksdorfii
brachyacantha
crj'thropoda
ularis
saligna
Fic. 1.CladogramofNorth-Americanblack-frtntedCrcituegicreatedbyPAUPusing38
morphologicalcharacters.
showninallanalysestobesisterspecies.Thetwounpublishedtreesonly
differinthelocationofC. suksdorfiiwhichiseitherbasal toC. cJo/zglasiiI
okennonii(publishedtree),basalto(B(Sa(R,E))),usingtheirepithetabbre-
viations,inoneunpublishedtreeorbasaltoboth(D,0)and(B(Sa(R,E))in
theother.
Theunionofallsevenblack-fruitedspecies insidethered-fruitedout-
groupsC.mollisandC.monogynadoesnot,however,indicatethattheblack-
fruitedgroupismonophyletic.Ananalysisusingallthered-fruitedspecies
wouldberequiredtogeneratesuchafindingandthatisnotthefunctionof
thisexercise.Rather,itistolocatethepositionofC.erythropoda,C.nvidaris,
C.salignaandC.brachyacantharelativetotheimmediateC.douglasiigroup.
ThecladogramendorsestheviewthatneitherC.salignanorC.rwularisare
partofthespeciesC. douglasii.Thefollowingkeysummarizesthediffer-
encesamongthefourspeciesinrwularisclade.
KEYTORIVULARIS-CAADV.OFBLACKANDBURGUNDY-FRUITEDCRATAEGUS
1.Stamens20;antlierscream;flowers10-12mmdiam.
2.Thorns1-1.5cmlong,decurved,grayish;twigsgrayish;overmatureflowers
orange-yellow;nutletswithlateralfacesplane C.brachyacantha
2.Thorns2-1cmlong,straight,black;twigscopper-colored;overmature
flowerswhitetodirtywhite;nutletswithlateralfacespitted C.saligna
1.Stamens10;antherspink;flowers15-17mmdiam.
.3.Leavesessentiallyunlobed,usuallymorethan2timesaslongasbroad,
widestnearthemiddle;fullyri|iefruitblack C.rivularis
3.Leavesevidentlyabout3-iobedperside;usuallyca. 1.5timesaslongas
broad,usuallywidestinthebasalthird;fullyripefruitusuallyburgundy
C.erythropoda
652 SiDA18(3)
TAXONOMICANDNOMENCLATHKALNO'I'I'SONINDIVIDUALSPFX;iES
CRATAEGUSERYTHROPODA—Chimicti^vviimon
Crataeguser>thr()j:KKlaAshe,NorthCarolinaAgric.Exp.Sta.Bull.175:113.1900.
Cnitaeg/mbuktri(irc-cne.nomennudum
Thisspecieshasbeendescribed,illustratedandmappedinmyrecentpaper
(Phipps 1999)whereitwasentered intoanewmonotypicseries,Cerrorm
J.B. Phipps,onaccountofitsdistinctnessfromothetred-ftuitedspecies.
Crataeguserythropodamaybebrieflycharacterizedbyitsovate-rhombic, +
glabrous,shallowly-lobed,smallishleaves;glabrousinflorescenceswith10-
stamenflowershavingpink-purpleanthers;suborbicularfruitwhichisburgundy
atmaturity(Ihaveneveronceseen'orange'asdescribedintheprotologue
byAshe(1900)andHolmgren(1997));shinycopperybarkon2-5cmthick
stemsandthornsmainly2-4cmlong.Crataeguserythropodaoccursinsites
withgroundwateravailable,ototherwisemesic,mainlyinintermountain
ColoradoandadjacentNewiVIexico,whereitisquitecommon.Holmgten
(1997)alsorecordsitfromeasternUtahandnortheasternArizona.The'or-
ange'fruitcolorcited mayrefertoimmaturefruitsincethetypewascol-
lected inAugust,but 1havenotmyselfseen thiscoloreven in mid-Au-
gust.Crataeguserythropodahaspresentednoseriousproblemsindelimitation
orrecognitionthoughitsrelationshipshavebeenhithertoobscure.Seethe
protologue(Ashe 1900)fotanotherfulldescription.Interestingly,Palmet
(1925)assignedC.erythopodatoser.Douglasianae,anideatakenupbyno-
oneelse,butneverthelesstheclosestapptoximationtodate.Observations
madeforthispaperindicatethatatanthesisC.erythropodaisalmostindis-
tinguishablefromC.nvidansinfloral/inflorescencecharacteristics.Significant
differencesinleafshapeandripefruitcolorexist,however,asindicatedin
thekey.
AspecimenofGteene'sftomtheLowerCimarronRiver,Colorado(NDG),
collectedin1896andlabelled'C.haken\anameneverpublished,isactu-
allyaperfectlyadequatespecimenofC.erythropodaAshe.
—
CRATAEGUSRIVULARIS Synonymy,CharacterizationandTypification
CrataegusrivularisNutt.ap.Torn&A.Gray,Fl.N.Amer. 1:464. 1840.
Cn/t,/c'ii//sdoii^^lctsiiLincll.var.rivulani(Niict.ap.Torr.&A.Gray)Sarg.,Gard.&
Forest i:81. 1902.
Aswith C. erythropoda, therehave notbeen serious problems in taxon
recognitionordelimitationalthoughsomefloristicauthorshavefollowed
Sargent(1889)inrelegatingC.nvularistovarietalrankunderC.douglasTi.
Crataegusnvularisisalocallycommon,evenlocallyabundant,speciesof
intermountainUSA,foundinmanylocationsinthisgenerallydryareawhere
thereisahighwater-table. I mapthiswidespreadspecies(Fig. 2)which
Phipps,Black-fruitedCrataegusfromthewesternUnitedStates 653
occursfromsouthernIdahotonorthernNewMexicoandfromsouth-cen-
tralUtaheasttotheRockyMountains.Itreaches6-7minfavorablesites
whereitmayformextensivethickets.Thebarkof2cmdiameterbranches
iscopper-coloredlikeC.erythropodaandthegenerallyfinethornsaremainly
2.5^cmlong.Thenormallyellipticleavesaretaperedateachend,subglabrous,
4—6-veinedandquiteunlobedonshort-shoots,beingbesetwithmanysharp,
forward-pointingfinemarginalteeth.Theglabrousinflorescencebearslarge
10-stamenedflowerswithpinkanthersandcanhardlybedifferentiatedfrom
thatofC. erythropoda. The numerous,oftenlarge, suborbicularfruitpass
throughastrikingdeepreddish-burgundycolortodeeppurpleandfinally
black.SeealsoSargent(1890)foradetailedcharacterizationandexcellent
platebyFaxon.IntermediateswithC.douglamarenotknownandC.rmdaris
isessentiallyallopatricwith thatmorenorthernspecieswhichfactsrein-
forcethefindingsofthecladisticanalysis.However,becauseWelsh(1982),
Holmgren(1997),andSargent(1889, 1890)haveincludedC. rmdarisin
C.douglasii,Iwouldliketoemphasizetheirdistinctionwiththefollowing
keycouplet.
1.Thornsfine,littlerecurved,usually2.5-5cmIcjng;barkoC1-2cmdiam.
branchesshinycopperybrown;flowers14—18mmdiameter,calyxlobes5—
8mmlong,long-attenuatetromabroadbase;leavesnarrowelliptic,unlobed
orrarelywith1—2verysmalllobesperside,taperedatbothends,acuteto
acuminateattheapex,thevenationsemi-camptodtomous;fruitsubglobose,
11—14mmdiam.,crimson-lakeripeningtoshinyblack C.rivularis
1.Tbrhaonrcnhsessttoaunt,toofgtreanyr-ebcruorwvne;d,fluoswuearlsly121.—51—53mcmmldoinagm;.,bacraklyoxfl1o—b2esc3m—4dimamm.
long,narrow-triangularfromabroadbase;leavesmuchbroader,mostcom-
monlybroadelliptictonarrow-obovateingeneralshape,usuallylobed(unlobed
inrarenarrow-leavedspecimens),thevenationclearlycraspedoclromousexcept
inrarenarrow-leavedspecimens,tisuallyobtusetosubacuteattheapex;fruit
generallyellipsoid,usually=S9mmdiameter,ripeningdullptirpletopurple-
blackorblack,generallywithratherstrongbloom C.douglasii,s.s.
Thesuiteofdifferencesissolarge,togetherwiththelackofintermediates
anddifferentdistribution,thatonecannotreasonablyplaceC rivularisin
thesamespeciesasC.douglasii.
InmakingC. rivularisavarietyofC. douglasiiSargent'soverallunder-
standingofthistaxonwaspoor,illustratedbyhisproviding(1890)anin-
accuraterangeextensionfromnorthwesternCaliforniatoPugetSoundand
hisstatement,"usuallyalowintricatelybranched shrub."Thisperhaps
helpstoexplainthecautiousrankchosenbySargent,anotorioussplitter
bymoderncriteria.
Crataegusrivularishasnotbeentypified,therefore,Ilectotypifyithere.
ThetypedescriptionforC,rivularis(Torrey&Gray1843)isofcharacteris-
ticbrevityandimprecisionfortheperiodandmightatfirstsightreferto
654 SiDA 18(3)
Fig.2.DistnbLirionolC..riviiLiyisNlitc.ap.TorrcyandA.Grayfromcollatedherbarium
retorcls.
C. cloii'^Lts'!}orevenC. Scdigrict^aswellasCothetaxon normallyassociated
withthename.
TheprotologueforCriviihiriscites"Orei;on,rivuletsintheRockyMountains"
asthetypeareawithNuttallascollector.Itishardtoknowexactlywhere
this is, becauseof theextensivenatureof theOregonTerritory in 1843.
Whatwecall C. nvi/laristodayisadistinct, well-collected taxonwitha
well-establishedrange(Fig. 2)which jnstreachessouthernIdaho,partof
whichisanacceptableareafor"RockyMountains."Putativesyntypema-
Phipps,Black-fruiteclCrataegusfromthewesternUnitedStates 655
terialsofC.rivulamfromBM,GHandPHwereborrowedinordertolectotypify
thisspecies.
OfthetwoPHspecimenssent(bothonthesamesheet)one(ontheright-
hand side)can berejecteddirectlyas it isaCanby-coUectedC. douglasii
from1(S73,whiletheleft-handspecmien,labelled'RockyMountains',isa
leafytwigwithoutreproductivematerialthatrequirescomparisonwiththe
protologue.BoththeBMandGHsheetshavespecimenscollectedbyNuttall
thatareareasonablematchforthesomewhatpoordescriptionintheprotologue
andareclearlythesamespeciesaseachother.Inviewoftheindifferentquality
oftheprotologueandtheconsequencethataspeciesotherthanwhatiscon-
ventionallycalledC.rivulansmighthavebeendescribed,IprovideinTable
2criticalcomparisonsbetweentheprotologueandputativetypes(cols. 1,
2),betweentheprotologueandC.doiiglas'ii(col.3)andbetweentheprotologue
andstandardinterpretationofC rividaris(col.4).
Severalpointsintheprotologuecannotbeassessedfromtheputativesyntypes
fromBMandGH. Thesearerhearborescentnatureoftheplant(notre-
cordedonthecollectionlabel)andfloweringcharacteristics(theputative
syntypesarepost-flowering).Also,Nuttallindescribingtheleavesas"ovate,
obovate"posessomethingofaproblem,asthesearenottermsthatwewould
applytodaytotheputativesyntypeleaves,whichareclearlynearertonar-
rowlyelliptic.Infact,inexammmgmanyprotologuesforC.seriesCoccineae
drawnupbyC.S.Sargentinthefirstandseconddecadesofthiscentury,I
haveobserved that"ovate"isalmostagenericleaf-shape.Inmyopinion,
havingalsoexaminedmanyotherCrataegustypedescriptionsfromthenineteenth
centuryandearlier,theterm "ovate"wasnotusedwith itspresentpreci-
sionandtherefore,withregardtoleaf-shape,theprotologuemaybesaidto
describethematerialoftheputativesyntypesinquestionwithadequate
accuracyfortheperiod.Theothercharactersmatchtheprotologue.Asto
thePHputativesyntype(col.2),thematchesbetweenitandtheprotologue
aresofewas to bealmostmeaningless. Moreover, thespecimen possibly
belongstoC.cnn-galli,aspreviouslystatedbyEgglestoninanannotation
onthesheet,butthisisaspeciesneverfoimdintheRockyMountainarea.
However,thespecimenalsoresemblesC.saligna,butbecauseitissterile,
conspecificityisuncertain.Thefinaltwocolumnsinthetablecomparethe
protologuewithtypicalC.douglasiiandwhatisnormallycalledC.rivularis
(alternativelyC.donglasitvar.rivularis).BecauseC.douglasuhaslobedleaf-
bladesand"short"thorns,whereasC.rivulanshaslongerthornsandunlobed
leaves,thelatterconstitutesthebettermatchfortheprotologue.Thus,the
choiceoflectotypeliesbetweeninadequatevegetativematerialofwhatis
possiblyC.salignaatPHandadequatespecimensclearlyrepresentingwhat
is normally called C. rivulansat BMand GH. Therefore, I lectotypify
656 ^"^11''^^1«('^)
!'abi,1'2.Comparisonofprorolo,i;iifcliaracteristicswithpnrativfsyntypcsofC.riniLirn(seclext)and
vichplantsofC.doiigLiMiandC.riviihivis.
matchfor matchfor
putative putative match match
syntypes syntype forC. forC.
protologuecharacteristics atBM,GH atPH doughisii rivularis
Aybonsci'Nt
NccirlygItihroNS +
Ndly,liiudiihir +
Lciiix-s -ovateorobovare ?,? ,+ V >
-obtuseorsometimesactire + +
-simplyorsomewhatnuisely + +
serrate
-attenuateintoashort]ienole + +
'I'biiyrn -"spines"loni^ +
Curymh-manyflowered ? ? ± ±
-glabrous ? ? us +
-flowerssmall ? ? + +
segmentsolealyxobtuseand ? + ±
veryshort
fnul -hluck
? = cannortlerermineamatch
+ = perfectmatch
± = reasonablemarch
us = tisuallyamarch
• - not.1matth
C. rivHlarhwiththeNuttallspecimen"RockyMountains"(holotypeBM;
isotypeGH)andretaintheusualinterpretationofthistaxon.
—
CRATAEGUSSALIGNA Synonymy,C:haractcrizationandTypification
CrataegussalignaCireene,Pittonia3:99- 1H96.
C>\itiieii,!/swhceleriA.Nelson,lint.Gaz.34:369. 1902.
Crutdnii^iischiiigliisiiLilull,var.cliichesueiLsisSA..Welsh,CireacBasinNattinilisr12:9.1982.
Crataegusudigmtisamedium-sizedtolargethicket-formingbushgrow-
ingalongstreamsandotherlocallymoistareasinintermountainColorado
andnortheastern Utah(Fig.3).Itsrangeisthemostrestrictedofthefour
speciesdealtwithinthispaper.InmanyrespectsitislikeC. riviiLirisbut
ithassmallerleaves, thorns,flowersandfruitand20creaminsteadol 10
pinkanthers.Thefruitismorefullyblack(darkerthanC. rivuhinswhen
sub-ripeinlateAugust)andtheleaf-teetharequitedifferentfromC.rividarn.
beingsmall,numerousandobtuse,ratherthanlongerandsharp.Moreover