Systematic wood anatomy of the Rosaceae

78 Pages·1992·7.6 MB·English

by  Zhang, Shu-Yin

BLUMEA 37 (1992) 81-158
Systematic wood anatomy of the Rosaceae
Shu-Yin Zhang
Rijksherbarium/Hortus Botanicus,Leiden,The Netherlands
Summary
Thewood anatomyofthe Rosaceae issurveyed andanalysed,basedonthe studyof280species
(c. 500 specimens)belongingto62 genera from differentpartsofthe world.Eighteen wood ana-
tomical charactershavebeenusedforapheneticandphylogeneticclassification. In thepheneticclas-
sification,12 groupsarerecognisedandcomparedwithHutchinson’s tribes. GroupsI-Vaccommo-
dateamixtureofrepresentatives fromSpiraeoideaeand Rosoideae genera (ortribes);GroupsVI-VII
comprise the Maloideae plusCercocarpus; GroupsVIII-IXarecomposedmainlyofthe tribe Quil-
lajeae,but includePrinsepia, andexclude Exochorda;GroupsX-XIIrepresentthe Prunoideae. All
woodytribes recognisedby Hutchinson encompass few closelysimilarwood anatomical groups ex-
cepttheCercocarpeae.TheQuillajeae,characterisedbypresumablyprimitive macromorphological
characters,appear tobequitederivedwood anatomically,and,accordingtothephylogeneticanalysis,
shouldbeincluded inthePrunoideae orberecognisedasaseparatesubfamily closetothePrunoideae.
Thewood anatomicalvariationpatterns supportabroad genericdelimitation inthe Maloideae,anda
narrow onein the Prunoideae. The wood anatomical evidence also suggests thatCercocarpus be
movedtothe Maloideae,and thatSorbariais the mostderived representativein theSpiraeoideae/
Rosoideae. Prinsepia, although distinct in its wood anatomyfrom thePrunus alliance,still forms
partofamonophyletic group togetherwiththe latter.Based onwood anatomical evidence,the sub-
division ofthefamily andtheoriginoftheMaloideae arediscussed.
WoodanatomicallytheRosaceae areisolatedwithin theorderRosales.Itishypothesisedthatthe
Rosaceae mighthave originatedin EastAsiainsteadofinthe tropics.
Individual genericdescriptions,a familydescriptionand atentative wood anatomical keytothe
generastudiedarealsopresented.
Introduction
ThewoodanatomyoftheRosaceaenativetoChinawas describedand discussed
in aprevious paper (Zhang & Baas, 1992). Thepresent paperextends theobserva-
tionto aworldwidesurvey, andaims ata contributiontoour understanding ofinter-
genericrelations andthephylogeny ofthefamily.
TheclassificationoftheRosaceaehaslong been in somedispute. Nowadays, the
Neuradeaeand theChrysobalaneae, formerly two tribesor subfamiliesofthe Rosa-
ceae(cf. Hutchinson, 1964; Yu, 1984), are usually treated as separate families:the
Neuradaceaeandthe Chrysobalanaceae (AiryShaw,1973;Cronquist, 1981; Schulze-
Menz, 1964;Stoffers, 1982; Takhtajan, 1980;Kalkman, 1988). In thepresent study,
theRosaceaeare delimitedinthe narrow sense.Foursubfamiliesare usually recog-
nisedin theRosaceae sensu stricto: Spiraeoideae, Rosoideae, Maloideae,andPrun-
oideae.Thefoursubfamiliesareratherdifferentin homogeneity, scopeandnatural-
ness, andareoften takenas unitsofdiscussion (Kalkman, 1988). For thesystematic
82 BLUMEA VOL. 37, No. 1, 1992
Table 1.EnumerationofgeneraoftheRosaceae.
GGeennuuss11 ((ssuubbffaammiillyy,, ttrriibbee))22 HHaabbiitt33 AApppprrooxxiimmaattee NNuummbbeerr ooffssppeecciieess
nnuummbbeerr ooffssppeecciieess ((ssaammpplleess))ssttuuddiieedd
AAccaaeennaa ((RR,,1144)) HHeerrbb cc.. 110000
AAddeennoossttoommaa ((RR,, 1166)) SShhrruubb 22 22 ((44))
AAggrriimmoonniiaa ((RR,, 1144)) HHeerrbb 1155
AAllcchheemmiillllaa ((RR,, 1144)) HHeeirbb 225500 11 ((11))
AAmmeellaanncchhiieerr ((MM,, 1188)) TTrreeee--SShhrruubb 66 33 ((33))
AAmmyyggddaalluuss ((PP,,66)) TTrreeee--SShhrruubb cc.. 4400 1100 ((2211))
AArreemmoonniiaa ((RR,, 1144)) HHeeribb 11
AArrmmeenniiaaccaa ((PP,,66)) TTrreeee cc..88 66 ((1177))
AArroonniiaa ((MM,, 1188)) SShhrruubb 22
AArruunnccuuss ((SS,,22)) HHeeribb 11
BBeennccoommiiaa ((RR,, 1144)) SShhrruubb 55
CCeerraassuuss ((PP,,66)) TTrreeee--SShhrruubb cc..110000 2200 ((4455))
CCeerrccooccaarrppuuss ((RR??,, 1155)) SShhrruubb--TTrreeee 66 22 ((44))
CChhaaeennoommeelleess ((MM,, 1188)) SShhrruubb 44 44 ((44))
CChhaammaaeebbaatiatia((RRoorrNN,, 1133)) SShhrruubb 22 11 ((11))
CChhaammaaeebbaattiaiarriaia((SS,,44)) SShhrruubb 22 11 ((11))
CChhaammaaeemmeelleess ((MM,, 1188)) SShhrruubb I1
CChhaammaaeerrhhooddooss ((RR,, 1122)) HHeerrbbss I1I1
CClliiffffoorrttiiaa ((RR,, 1144)) SShhrruubb 111155
CCoolleeooggyynnee ((RR??,, 1155)) SShhrruubb 11 11 ((11))
CCoolluurriiaa ((RRoorrNN,, 1133)) HHeerrbb 66
CCoommaarreellllaa ((RR,, 1122)) HHeeribb 22
CCoottoonneeaasstteerr ((MM,, 1188)) SShhrruubb--TTrreeee cc.. 5500 66 ((88))
CCoowwaanniiaa ((RR oorrNN,, 1133)) SShhrruubb 33--44 11 ((11))
CCrraattaaeegguuss ((MM,, 1188)) SShhrruubb--TTrreeee cc.. 228800 1188 ((2233))
CCyyddoonniiaa ((MM,, 1188)) TTrreeee 11 11 ((33))
DDiicchhoottoommaanntthheess ((MMoorrNN,, 1188)) TTrreeee 11 11 ((11))
DDooccyynniiaa ((MM,, 1188)) TTrreeee 55 22 ((33))
DDrryyaass ((RRoorrNN,, 1133)) SShhrruubb 33--44 11 ((22))
DDrryymmooccaalllliiss ((RR,, 1122)) HHeerrbb cc..3300
EErriioobboottrryyaa ((MM,, 1188)) TTrreeee--SShhrruubb 2277 44 ((1100))
EErriioolloobbuuss ((MM,, 1188)) TTrreeee 11 11 ((11))
EExxoocchhoorrddaa ((SSoorrPP,,11)) SShhrruubb 44 11 ((11))
FFaalllluuggiiaa ((RR oorrNN,, 1133)) SShhrruubb 11 11 ((11))
FFiilliippeenndduullaa ((RR,,77)) HHeerrbb 1100
FFrraaggaarriiaa ((RR,, 1122)) HHeerrbb cc.. 1122
GGeeuumm ((RRoorrNN,, 1133)) HHeerrbb 665555
HHaaggeenniiaa((RR,, 1144)) TTrreeee 22 11 ((22))
HHeessppeerroommeelleess ((MM,, 1188)) SShhrruubb 1155 22 ((22))
HHeetteerroommeelleess ((MM,, 1188)) SShhrruubb 11 11 ((44))
HHoollooddiissccuuss ((SS,,99)) SShhrruubb 88 11 ((22))
HHoorrkkeelliiaa ((RR,, 1122)) HHeerrbb 1177
HHoorrkkeelliieellllaa ((RR,, 1122)) HHeeribb 33
IIvveessiiaa ((RR,, 1122)) HHeeribb 2233
KKaaggeenneecckkiiaa ((SSoorrNN,, 11)) TTrreeee 33 11 ((11))
KKeellsseeyyaa ((SS,,22)) SShhrruubb 11
KKeerrrriiaa ((SS,,RRoorrNN,, 1111)) SShhrruubb 11 11 ((22))
LLaauurroocceerraassuuss ((PP,,66)) TTrreeee--SShhrruubb cc..8800 99 ((2288))
LLeeuuccoossiiddeeaa ((RR,, 1144)) SShhrruubb 11
Shu-Yin Zhang:Systematic wood anatomy ofthe Rosaceae 83
(Table 1 continued)
GGeennuuss11 ((ssuubbffaammiillyy,,ttrriibbee))22 HHaabbiitt33 AApppprrooxxiimmaattee NNuummbbeerr ooffssppeecciieess
nnuummbbeerr ooffssppeecciieess ((ssaammpplleess)) ssttuuddiieedd
LLiinnddlleeyyaa ((SSoorrMM,, 11)) TTrreeee 22 11 ((11))
LLuueettkkeeaa ((SS,,22)) SShhrruubb--HHeerrbb 11
LLyyoonnootthhaammnnuuss ((SS,, 11)) TTrreeee--SShhrruubb 11 11 ((11))
MMaaddddeenniiaa ((PP,,66)) TTrreeee 44
MMaallaaccoommeelleess ((MM,, 1188)) SShhrruubb 22
MMaalluuss ((MM,, 1188)) TTrreeee--SShhrruubb cc.. 2255 1166 ((2266))
MMaarrggyyrriiccaarrppuuss ((RR,, 1144)) SShhrruubb 11
MMeessppiilluuss ((MM,, 1188)) TTrreeee 22 11 ((22))
MMiiccrroommeelleess ((MM,, 1188)) TTrreeee--SShhrruubb cc.. 1155 66 ((1177))
NNeeiilllliaia ((SS,,33)) SShhrruubb 1111
NNeevviiuussiiaa ((SS,, RRoorrNN,, 1111)) SShhrruubb 11
OOrrtthhuurruuss ((RR oorrNN,, 1133)) HHeerrbb 22
OOssmmaarroonniiaa ((PPoorrNN,,55)) TTrreeee--SShhrruubb 11--22 11 ((22))
OOsstteeoommeelleess ((MM,, 1188)) SShhrruubb 44--55 11 C(1D)
PPaadduuss ((PP,,66)) TTrreeee--SShhrruubb cc..2200 1133 ((3311))
PPeerraapphhyylllluumm ((MM,, 1188)) SShhrruubb 11 11 ((11))
PPeettrroopphhyyttuumm ((SS,,22)) SShhrruubb 55 11 ((11))
PPhhoottiinniiaa ((MM,, 1188)) SShhrruubb--TTrreeee 4400 99 ((2222))
PPhhyyssooccaarrppuuss ((SS,,33)) SShhrruubb 1100 11 ((22))
PPllaaggiioossppeerrmmuumm ((PP,,66)) SShhrruubb 22
PPoollyylleeppiiss ((RR,, 1144)) SShhrruubb--TTrreeee 1155 33 ((33))
PPoorrtteerraanntthhuuss ((SS,,44)) HHeerrbb 22
PPoottaanniinniiaa ((RR,, 1155)) SShhrruubb 11
PPootteennttiillllaa ((RR,, 1122)) HHeerrbb--SShhrruubb 55000055 11 ((22))
PPootteerriiddiiuumm ((RR,, 1144)) HHeerrbb 22
PPootteerriiuumm((RR,, 1144)) HHeerrbb cc..2255
PPrriinnsseeppiiaa ((PPoorrNN,,66)) SShhrruubb 33--44 22 ((22))
PPrruunnuuss ss.. ssttrr.. ((PP,,66)) TTrreeee--SShhrruubb cc.. 3300 99 ((1188))
PPsseeuuddooccyyddoonniiaa ((MM,, 1188)) SShhrruubb--TTrreeee 11
PPuurrppuussiiaa ((RR,, 1122)) HHeerrbb 11
PPuwrsshhiiaa ((RRoorrNN,, 1133)) SShhrruubb 22 22 ((22))
PPyyggeeuumm ((PP,,66)) TTrreeee--SShhrruubb cc.. 4400 1166 ((2233))
PPyyrraaccaanntthhaa ((MM,, 1188)) SShhrruubb cc..2200 11 ((11))
PPyyrruuss ((MM,, 1188)) TTrreeee--SShhrruubb cc..2200 1177 ((3333))
QQuuiillllaajjaa ((SS oorrNN,, 11)) SShhrruubb--TTrreeee 44 11 ((11))
RRaapphhiioolleeppiiss ((MM,, 1188)) TTrreeee--SShhrruubb 1144 44 ((88))
RRhhooddoottyyppooss ((RR,, 1100)) SShhrruubb 11
RRoossaa ((RR,, 1177)) SShhrruubb cc.. 110000 1155 ((2200))
RRuubbuuss ((RR,, 88)) SShhrruubb cc.. 225500 99 ((1111))
SSaanngguuiissoorrbbaa ((RR,, 1144)) HHeerrbb cc.. 1188
SSaarrccooppootteerriiuumm ((RR,, 1144)) SShhrruubb 11 11 ((33))
SSiibbbbaallddiiaa((RR,, 1122)) HHeerrbb 88 11 ((11))
SSiibbiirraaeeaa ((SS,,22)) SShhrruubb 22 I1 ((22))
SSoorrbbaarriiaa ((SS,,44)) SShhrruubb cc..99 44 ((44))
SSoorrbbuuss ((MM,, 1188)) TTrreeee--SShhrruubb 8855 2233 ((3344))
SSppeenncceerriiaa ((RR,, 1144)) HHeerrbb 22
SSppiirraaeeaa ((SS,,22)) SShhrruubb 7700 I1I1 ((1155))
SSppiirraaeeaanntthhuuss ((SS,, 1144)) SShhrruubb 11
SStteepphhaannaannddrraa ((SS,,33)) SShhrruubb 44 11 ((22))
84 BLUMEA VOL. 37, No. 1, 1992
(Table 1continued)
GGeennuuss11 ((ssuubbffaammiillyy,, ttrriibbee))22 HHaabbiitt33 AApppprrooxxiimmaattee NNuummbbeerr ooffssppeecciieess
nnuummbbeerr ooffssppeecciieess ((ssaammpplleess)) ssttuuddiieedd
SSttrraannvvaaeessiiaa ((MM,, 1188)) TTrreeee 55 11 ((11))
TTeettrraagglloocchhiinn ((RR,, 1144)) SShhrruubb 88
VVaauuqquueelliinniiaa((SSoorr MM,,11)) TTrreeee--SShhrruubb 1100 11 (01))
WWaallddsstteeiinniiaa ((RRoorrNN,, 1133)) HHeerrbb 66
TToottaallss:: 110022 OOvveerr33//44aarree OOvveerr22775500ssppeecciieess 228800((449911))
wwooooddyyggeenneerraa
1) The genericconceptsaremainly thoseadoptedby Kalkman (1988). However,Prunuss.1. (in
the senseofKalkman)ispresentedhere asseven smaller genera (seeZhang& Baas,1992)fol-
lowing Yu (1986):Amygdalus,Armeniaca, Cerasus,Laurocerasus, Padus,Prunus s.str.,and
Pygeum.The sameappliestoPhotinia whichexcludesStranvaesiaandHeteromeles here.
2) Followingsubfamily andtribeassignmentsaccordingtoKalkman (1988)andHutchinson (1964),
respectively.
— M:Maloideae;P:Prunoideae;R:Rosoideae;S:Spiraeoideae;N: Newseparatesubfamily, as
proposedbydifferentauthors,butnotgenerallyaccepted.
— 1=Quillajeae;2=Spiraeeae; 3=Neillieae;4=Gillenieae;5=Osmaronieae;6=Pruneae;
7=Ulmarieae;8=Rubeae;9=Holodisceae; 10=Rhodotypeae; 11=Kerrieae;12=Poten-
tilleae; 13=Dryadeae; 14=Poterieae; 15=Cercocarpeae; 16=Adenostomeae;17=Roseae;
18=Maleae (orPomeae).
3) Plant habitsaregivenhere mainlyaccording toHutchinson (1964)andYu (1986;forPrunuss.1.
only).
4) Theapproximatenumber ofspeciesforthegeneralisted aboveis basedonMabberley(1987)and
Yu(1986;only forPrunuss.l.).
5) Potentilla andGeuminclude somesegregates,accordingtoKalkman (1988).
arrangement,delimitationofsubfamiliesand genera,andthenomenclaturepresented
in this study, seeKalkman(1988). However,Photinias.l.(cf. Kalkman, 1973) and
Prunus s.1.(cf.Kalkman, 1965)are presented hereassmalleralliances(seethenotes
to Table 1). The only purpose for such atreatment in thepresent survey is to see
whetherthereis any wood anatomicalevidenceforsmaller genera.Foreasy refer-
ence, thegeneraoftheRosaceae,theirapproximate total numberofspecies, andthe
numberofspecies and samples studied are presented in Table 1. In addition, itis
also indicatedto whichtribesandfamiliestheindividualgenera(probably) belong.
As showninTable 1, 62generaincluding afew(sub)herbaceous generalikeAlche-
milla, Potentillaand Sibiraea,were surveyed in thepresent study, covering 16out
of 18 tribesrecognised by Hutchinson (1964). Only theherbaceoustribeUlmarieae
(consisting only ofFilipendula) andthewoody tribeRhodotypeae (Rhodotypos on-
ly) were notincluded.
Early studieson thewood anatomyofRosaceae were summarisedby Solereder
(1899 & 1908) andMetcalfe& Chalk(1950). For an updated literatureliston wood
anatomyofthefamily, seeZhang & Baas(1992).
Shu-Yin Zhang:Systematic wood anatomy of the Rosaceae 85
MATERIALS AND METHODS
Materials
Intotalc.500specimens were studied.Materialwas obtainedfromvariousinsti-
tutionalwood collections (see Acknowledgements). Many samples includedin the
presentsurvey are unvouchered.However, nomenclature was brought up to date
withrecent taxonomicrevisions, supposing thattheoriginal identificationhadbeen
correct. Synonyms are given ifnecessary. For the microtechnicalprocedures and
descriptive conventions followed, see Zhang & Baas (1992). The observationson
Chinesespecies are incorporated in variousgeneric descriptions given here. Ifstud-
iesofmaterialfromoutsideChina hardly modifiedourearliergeneric descriptions
thereaderisreferredtotheearlierpaper(Zhang & Baas, 1992).
Cladistic analysis
ThecomputerprogrammeHENNIG86 was used inthepresent study forthecla-
distic analysis ofwood anatomicalcharacters ofthe Rosaceae (Farris, 1989;Van
Welzen, 1989).
HENNIG86 uses the Wagner algorithm combinedwith parsimony (cladogram
withtheminimumnumberofcharacterstatechanges, = steps,willbeselectedas the
best hypothesisofthephylogeny). HENNIG86 was mainlyused withunorderedchar-
acters by givingthecommandCCODE-.;. As tree calculating algorithm MHENNIG,
in combinationwithBB; was used.MHENNIG onlycalculatesseveraltrees, ofwhich
themostparsimonious ones willbe stored. Thesedonot haveto bethe mostparsi-
monious ones amongalltrees by necessity, nor the only mostparsimonious trees.
BB (branch and bound) uses theMHENNIG trees and usually finds, by modifying
branches, all most parsimonious trees (the option IE insteadof thecombination
MHENNIG-BB is sure to findall mostparsimonious trees, butwhenthedatamatrix
is complex andcontainsmuchhomoplasy therunning timeofoption IEbecomestoo
long, days insteadofhourswithMHENNIG-BB).
TheWagner algorithm needsanapriori designated outgroupofwhichthecharac-
terstatesareregarded as plesiomorphic (primitive). Foreach charactertheoutgroup
always (ifno question markisplaced inthe datamatrix) assigns one characterstateas
thestarting pointofatransformationseries.
Ifmore than one parsimonious tree was foundcharacter weighting was applied
withthecommand: XSTEPS W; followedby the branchand boundcommandBB;.
Thecharacterweights are automatically calculatedby multiplying theconsistency
index withthe retentionindex (Farris, 1989) and scaling these from0to 10. This
procedure isrepeated untilone cladogramremainsor untilthecharacterweights do
not change anymore.
WOOD ANATOMICAL FAMILY DESCRIPTION
Inaprevious paper(Zhang &Baas, 1992),anelaboratesurvey ofthewoodana-
tomicalvariation in ChineseRosaceae was given. Fromthe study ofa substantial
numberofadditionalgeneraandspecies fromoutsideChinait appeared that,except
for someminorextensionsofquantitative ranges, thisearlier survey almostcover-
86 BLUMEA VOL. 37, No. 1, 1992
ed the entirerange present in the family. As a consequence,the wood anatomical
description ofthe entirefamily given below hardly differsfromthe one given for
Chineserepresentatives only. Where appropriate, notes (numbers between square
brackets) are given ontheextendedrange ofsomecharacters.Seealso Tables2&6,
theindividualgeneric descriptions,andFigures 1-16.
Growth rings moreorless distinctin most genera[1],rarely faintor absent,the
boundariesinfrequently wavy,mostly markedby rows ofradially flattenedlatewood
fibres, sometimesby differencesinvessel diameteras wellas vessel frequencybe-
tween latewoodand subsequent earlywood,orrarely by marginal parenchyma bands
(Figs. 1-8). Wooddiffuse-porous, semi-ring-porous orring-porous.
Vessels 40-300(5-600)/sq.mm, 4-99% solitary [2], remainder inoblique,
tangential andradial multiplesof2-4(-12), vessels insome taxa inanoblique,radial
or dendriticpattern [3], round,oval to angular, tangential diametermostly 20-80
(16-175) pm,radialdiameter30-100(20-220) pm,walls 1—3(—5) pmthick (Figs.
1-8). Vessel elementlength 150-700 (130-930) pm, length/tangential diameter
(L/D)ratio3-30. Perforationsexclusively oralmostexclusively simple inoblique
toalmost horizontalendwalls; sporadic scalariformand/or irregular perforations
notedinsomespecies. Intervesselpits nonvestured(except inSpiraea p.p.), usually
alternate,round,oval topolygonal, 2-12(-20) pmin diameter,with slit-like, occa-
sionally oval tocoalescentapertures.Vessel-ray and vessel-parenchyma pits mostly
LegendstoFigures1—16:
Figs. 1—4.TS,x43. — 1:Growthringboundariesabsent andvessels oftwo slightlydifferent size
classes inHageniaabyssinica.— 2:Growth ringboundaries faintto absentand vessels diffuse in
Polylepisincana. —3:Growthringboundaries markedby 1—3-seriatemarginalparenchymabands
—and traumatic gum ductpresentinLaurocerasus hypotricha. 4:Growth ringboundaries distinct,
marked by rows ofradiallyflattened latewood fibres;vessels in aweaklyobliquepatternand witha
highdegreeofgroupingin Laurocerasus spinulosa.
Figs. 5—7. TS,x 43. — 5: Growthringboundaries faint,marked by marginalparenchyma band;
vessels moderatelylarge inPygeum topengii.—6:Growth ring boundaries distinct and weakly
wavy, markedby rows ofradiallyflattenedlatewood fibres, vesselsin aweaklyobliquepatternand
with ahighdegreeofgroupingin Sarcopoteriumspinosum.—7; Woodsemi-ring-porous,vessels
mostly solitaryand round to oval in outline,and parenchymarelatively abundant in Cercocarpus
betuloides.F— ig. 8.TS,x 116.Wood ring-poroustosemi-ring-porous,vessel outline angularin
Potentilla fruticosa.
Fig. 9.TLS,x 116.Vasculartracheidspresentin Sarcopoteriumspinosum.—Fig. 10.TLS,x576.
One toseveralprismaticor fragmentedcrystalspernon-enlargedaxialparenchymachamberin Spi-
raea chamaedryfolia.—Fig. 11.RLS,x 43. Rays composed exclusively ofprocumbentcells in
Hageniaabyssinica. —Fig. 12.TLS,x 116.Rays 1(—2)-seriateinPotentilla fruticosa.
F— ig. 13. TLS,x 11F6.Rays 1—4-seriate,but reliatively shgort in VauqueliniaCalifor.nia. 14.
TLS,x 116.Rays 1—3-seriate,butquitetall inKageneckia lanceolata.—Fig. 15. TLS,x43. Rays
oftwo distinctsizes in Hageniaabyssinica.—Fig. 16.TLS,x 288. Juvenilisticrays inStephanan-
dra chinensis.
Shu-Yin Zhang:Systematic wood anatomy ofthe Rosaceae 87
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