Notes on taxonomy and pollen of Malesian Daphniphyllum (Daphniphyllaceae)

14 Pages·1996·2.2 MB·English

by  Huang, Tseng-Chieng

BLUMEA 41 (1996) 231-244
Notes on taxonomyandpollenof
Malesian Daphniphyllum(Daphniphyllaceae)
Tseng-Chieng Huang
DepartmentofBotany,National TaiwanUniversity, Taipei 106,Taiwan,Republic ofChina
Summary
This paper is a precursor tothe revision ofthe familyDaphniphyllaceaein Flora Malesiana,in
which 16species ofthe genusDaphniphyllumwill betreated. Inthepresentpaperadescription is
givenofonenewvariety,D. gracileGagevar.newirelandum, andfive newcombinations havebeen
made,D. ceramense, D. dichotomum,D.papuanum Hallier f.var. tuberculatum, D. sumatraense,
andD. timorianum. Anenumeration ofthe 16species aswill betreated inFlora Malesiana is given
here;the numberingofthenew variety and the five newcombinations in thepresent paperis the
sameaswill beused in Flora Malesiana. A keytosections,subsections and series is given. Pollen
features,adistribution map ofthetwo sections,and aline drawingofthenew varietyareprovided.
Introduction
Forming a monotypic family,Daphniphyllaceae, thegenusDaphniphyllum consists
ofabout 30species, with 16ofthem intheMalesianregion. The species are distrib-
utedfromIndiatoJapan andfromCentralChinatoNewGuinea(Fig. 1).They grow
onhillsides, burntareas, grasslands ormontane forests ataltitudesof300-4000m.
Monographic studyofDaphniphyllum was initiatedby MiillerArgoviensis (1869),
and followedby Rosenthal(1919) and by T.C. Huang (1965, 1966). Six species
werereported fromtheMalesianregion inmy treatment (Huang, 1966);three sec-
tionswerealsoproposed: sectionDaphniphyllum [including D. glaucescens Blume,
D. gracile Gage, D. philippinense Huang and D. woodsonianumHuang], section
Lunata Huang [including D. laurinum(Benth.) Baill.],andsection StaminodiaHuru-
sawa [including D.parvifolium Quisumb. & Merr.]. Sincethen, norevision nor
monographic treatment hasbeendonefortheMalesianregion.
Morphological featuresusedalonecannotbeexpected toreach theidealclassifica-
tionofthegenusDaphniphyllum. Iaccept moreoftheolderspecific names forthis
paperthan inmy previous work.Morefieldandexperimental workis neededbefore
bettertaxonomicresults can be obtained,becausethese plants possess simple fea-
turesandregional variationamongtheislands.Therefore, thefollowing taxonomic
treatmentis stillaprovisional one.
Daphniphyllaceae have beenplacedinvariousorders(Huang, 1965,1966). There
aresomerecenttreatments,such asinPittosporales by Thome(1976,1983). Hutchin-
son (1973) and Takhtajan (1980) placed itinHamamelidales.Dahlgren (1983) and
Thorne(1992) consideredthefamily asamemberofBalanopales (Buxales). Sutton
(1989) andZhang & Lu(1989) placeditin Daphniphyllales with closestrelationship
toHamamelidalesas suggested by Huang (1965) andCronquist (1983).
232 BLUMEA —Vol. 41, No. 1, 1996
POLLEN MORPHOLOGY
Thenew dataobtainedfromascanning electronmicroscopic study showthatpollen
grains are stenopalynous, isopolar, 3(-4)-aperturate. Grain size is 11-18pm.Grain
shape is subprolate tosuboblate(P/E= 1.21-0.85).Theequatorial outlineiscircular
tosemi-circular. Theapertureis colpateto colporoidate. Exineornamentsare irregu-
larlyperforate in sectionLunata(D. laurinum) and fossulateinsectionDaphniphyl-
lum (Table 1; Plates 1-3).
Table 1. Pollen data of Daphniphyllum.
TTaaxxaa OOrrnnaammeennttss PP//EE AAmmbb VVoouucchheerrss
((uH.mm))
SSeeccttiioonnLLuunnaattaa
DD.. llaauurriinnuumm iirrrreegguullaarrllyy 1111--1122..55//1111-1-111.5.5 1111--1122 SSooeeppaaddmmoo&&MMaahhmmuudd
ppeerrffoorraattee pprroollaattee--sspphhaaeerrooiiddaallttoo 11119933
oobbllaattee--sspphhaaeerrooiiddaall DDiinnggHHoouu 775588
WWhhiitlmmoorree 33778899
SSeeccttiioonnDDaapphhnniipphhyylllluumm
DD.. bboorrnneeeennssiiss ffoossssuullaattee 1133..55--1155..44//1155.8.8--1166..22 1144--1155 MMeeiijjeerr-5544227755
ssuubboobbllaattee ttoooobbllaattee-- HHaavvilialanndd11007700
sspphhaaeerrooiiddaall
DD.. ggrraacciillee ffoossssuullaattee 1133..99--1155..88//1144.2.2--1177..77 1177--1188 KKaallkkmmaann 44999944
oobbllaattee--sspphhaaeerrooiiddaallttoo FFrrooddiinn 2266882299
ssuubboobbllaattee HHaarrttlleeyy1122774488
SStteevveennss && VVeellddkkaammpp
5544223300
DD.. ppaappuuaannuumm ffoossssuullaattee 1122..88--1177..55//1111.9.9--1177..66 1144--1199 FFrrooddiinn 2266229911
vvaarr..ttuubbeerrccuullaattuumm oobbllaattee--sspphhaaeerrooiiddaallttoo JJaaccoobbss99441144
pprroollaattee--sspphhaaeerrooiiddaall FFlleennlleeyy22661111
BBrraassss3311446600
DD.. ttiimmoorriiaannuumm ffoossssuullaattee 1122..99--1144..88//1122.2.2--1144..55 1133--1177 KKoosstteerrmmaannss&& WWiirraawwaann
ssuubbpprroollaatteettoo oobbllaattee-- 448844
sspphhaaeerrooiiddaall MMeettzmneerr225533,, 225555
SScchhmmuuttzz 33448844
Plates 1—3.Illustrations ofpollengrainsofDaphniphyllum.White scale bars= 3µm; black scale
bars = 1 µm, respectively. Photos 1,5,9, 13and 17areequatorial views;2,6, 14, 18and 20 are
polarviews;3,7, 11, 15and 19aremesocolpiumareas;4, 8, 12, 16 and20 are apocolpiumareas.
Photos 1—4 show irregularly perforate ornamentation and 5—20 show fossulateornamentation. —
1—4: Daphniphyllumlaurinum (Benth.) Baill. (Soepadmo& Mahmud 1193);5—8: D. papuanum
Hallier f.(Frodin2629);9—12: D. gracileGage(Kalkman4994); 13—16:D.bomeenseStapf(Meijer
54275); 17—20:D. timorianum (T.C.Huang)T.C.Huang(Metzner253).
T.C.Huang: Malesian Daphniphyllum 233\1
Forlegends,seepage 232.
234\2 BLUMEA —Vol. 41, No. 1, 1996
Forlegends,seepage 232.
T.C.Huang: Malesian Daphniphyllum 235\3
Forlegends,seepage 232.
Fig. 1.Distribution map ofDaphniphyllumsectionDaphniphyllum(—————) and section Lunata
( ......).
236 BLUMEA —Vol. 41, No. 1, 1996
SYSTEMATIC TREATMENT(Table2)
Key to sections, subsections and series
la. Anthersusually oblongor elliptic, compressed or oblique, extrorse, completely
free, with a usually long filament; calyx present, usually caducous, atleast in
pistillateflowers, or absent; drupes smoothto tuberculate, usually not glaucous;
pollen larger than 12.7by 12.7pm;exinefossulate
1. SectionDaphniphyllum 2
b. Antherslunate, introrse, coherentatapex, subsessile; calyx persistent, atleastin
staminate flowers; drupes strongly tuberculate, usually distinctly glaucous;pol-
lengrains smallerthan 12.7by 12.7pm; exineirregularly perforate
2.SectionLunata
2a. Calyx4-6-lobed;drupes fromlustrousand smoothto verrucose ortuberculate;
leafblades usuallypapillate, chartaceoustocoriaceous, allveinsprominentand
reticulateabove 1A.SubsectionDaphniphyllum 3
b. Calyx absentorrarely 1-or2-lobed, linear, affixed tobaseofstamens; drupes
usually tuberculate; bladesusually epapillate, membranousto chartaceous, all
veinsobscureanddelicateor impressed above IB. SubsectionStaminodia
...
(D. parvifolium)
3a. Calyx present inbothstaminateandpistillate flowers 4
b. Calyx presentinpistillate flowers only lAc. SeriesUnicalycifera
(D. buchananiifolium, celebense, woodsonianum)
4a. Calyx inarticulate, usually persistent in staminateflowersand shorterthan the
androeciumand gynoecium lAa. SeriesDaphniphyllum
(D. borneense, ceramense, dichotomum,glaucescens, lancifolium,
luzonense, scortechinii, sumatraense,timorianum)
b. Calyx articulate, caducous and usually longer thantheandroeciumand gynoe-
cium lAb.SeriesLongicalycifera
(D. gracile,papuanum)
Enumerationof Malesian species 1
Daphniphyllum. Blume,Bijdr. (1826) 1153.
1. D.borneense Stapf 8. D. lancifolium Hook. f.
2. D.buchananiifoliumHallier f. 9. D.laurinum (Benth.)Baill.
3. D.celebense K.Rosenthal 10. D.luzonenseElmer
4. D. ceramense (T.C.Huang)T.C.Huang 11. D.papuanum Hallier f.
5. D. dichotomum(T.C.Huang)T.C.Huang a. var.papuanum
6. D. glaucescensBlume b. var.tuberculatum(T.C.Huang)
a.var.glaucescens T.C. Huang
b.blumeanum (Baill.exMull. Arg.) 12. D.parvifolium Quisumb.& Merr.
J.J. Sm. 13. D.scortechinii Hook.f.
7. D.gracileGage 14. D.sumatraense (T.C.Huang)T.C.Huang
a.var.gracile 15. D.timorianum (T.C. Huang) T.C.Huang
b. var.newirelandumT.C. Huang 16. D. woodsonianum T.C.Huang
1) Speciesto be treatedin Flora Malesiana;namesprintedin bold typehave been treated in the
presentpaper.
T.C.Huang: Malesian Daphniphyllum 237
Table2. Comparison of the Malesian taxa ofDaphniphyllum accepted by me in the
past (1965/66)and in thepresenttreatment.
SSeeccttiioonn 11..LLuunnaattaaHHuuaanngg SSeeccttiioonn22..LLuunnaattaaHHuuaanngg
11. DD..llaauurriimnuimm ((BBeenntthh..))BBaaiillll.. 99.. DD.. llaauurriinnuumm ((BBeenntthh..))BBaaiillll..
SSeeccttiioonn 22..DDaapphhnniipphhyylllluumm BBlluummee SSeeccttiioonn 11..DDaapphhnniipphhyylllluumm BBlluummee
SSuubbsseeccttiioonn11..DDaapphhnniipphhyylllluumm
SSuubbsseeccttiioonn11.. LLoonnggiiccaallyycciiffeerraa HHuuaanngg SSeerriieess 22..LLoonnggiiccaallyycciiffeerraa ((HHuuaanngg))HHuuaanngg
22 aa..DD.. ggrraacciilleeGGaaggee 77.. DD.. ggrraacciilleeGGaaggee
77 aa..vvaarr..ggrraacciillee
77 bb.. vvaarr..nneewwiirreellaanndduumm HHuuaanngg
1111.. DD..ppaappuuaannuumm HHaalllliieerr ff..
1111 aa..vvaarr..ppaappuuaannuumm
22 bb.. vvaarr..ttuubbeerrccuullaattuumm HHuuaanngg 1111 bb.. vvaarr..ttuubbeerrccuullaattuumm ((HHuuaanngg))HHuuaanngg
SSuubbsseeccttiioonn22..DDaapphhnniipphhyylllluumm SSeerriieess 11.. DDaapphhnniipphhyylllluumm
33.. DD..ggllaauucceesscceennssBBlluummee
33 aa..ssuubbsspp.. ggllaauucceesscceennss
33 aaaa..vvaarr..ggllaauucceesscceennss
33 aabb.. vvaarr..bblluummeeaannuumm ((BBaaiillll..eexx
MMuullll..AArrgg..)) JJ..JJ..SSmm..
33 bb.. ssuubbsspp..bboorrnneeeennssee ((SSttaappff)) HHuuaanngg 11.. DD..bboorrnneeeennsseelSSttaappff
33 cc..ssuubbsspp..cceerraammeennsseeHHuuaanngg 44.. DD..cceerraammeennsseeI((HHuuaanngg))HHuuaanngg
33 dd..ssuubbsspp.. cceelleebbeennssee ((KK..RRoosseenntthh..))HHuuaanngg ((-->> SSuubbsseeccttiioonn 22ssppeecciieess 33))
33 ee..ssuubbsspp.. ddiicchhoottoommuummHHuuaanngg 55.. DD.. ddiicchhoottoommuumm ((HHuuaanngg))HHuuaanngg
66.. DD..ggllaauucceesscceennssBBlluummee
66 aa..vvaarr..ggllaauucceesscceennss
66 bb.. vvaarr..bblluummeeaannuumm ((BBaaiillll..eexx
MMuullll..AArrgg..)) JJ..JJ..SSmm..
33 ff..ssuubbsspp..illaanncciiffoolliiuumm ((HHooookk,,ff..))HHuuaanngg 88.. DD..llaanncciiffoolliiuummHHooookk.. ff..
33 gg..ssuubbsspp.. lluuzzoonneennsseeEEllmmeerr 1100.. DD..lluuzzoonneennsseeEEllmmeerr
33 hh..ssuubbsspp.. ssccoorrtteecchhiinniiii HHooookk.. ff.. 1133.. DD..ssccoorrtteecchhiinniiii HHooookk.. ff..
33 ii.. ssuubbsspp..,ssuummaattrraaeennssee HHuuaanngg 1144.. DD.. ssuummaattrraaeennssee ((HHuuaanngg))HHuuaanngg
33 jj..ssuubbsspp.. ttiimmoorriiaannuumm HHuuaanngg 1155.. DD..ttiimmoorriiaannuumm((HHuuaanngg))HHuuaanngg
33 kk.. ssuubbsspp..bbuucchhaannaanniiiiffoolliiuumm((HHaalllliieerrff..))
HHuuaanngg ((——»» SSeerriieess 33,,ssppeecciieess22))
SSuubbsseeccttiioonn33..UUnniiccaallyycciiffeerraa HHuuaanngg SSeerriieess 33..UUnniiccaallyycciiffeerraa ((HHuuaanngg))HHuuaanngg
44.. DD.. pphhiilliippppiinneennsseeHHuuaanngg 22.. DD..bbuucchhaannaanniiiiffoolliiuummHHaalllliieerr ff..
55.. DD.. wwooooddssoonniiaannuumm HHuuaanngg 1166.. DD.. wwooooddssoonniiaannuumm HHuuaanngg
SSeeccttiioonn 33..SSttaammiinnooddiiaaHHuurruussaawwaa SSuubbsseeccttiioonn22..SSttaammiinnooddiiaa((HHuurruussaawwaa))HHuuaanngg
33.. DD..cceelleebbeennsseeKK..RRoosseenntthhaall
66.. DD.. ppaarrvviiffoolliiuummQQuuiissuummbb..&& MMeerrrr.. 1122.. DD..ppaarrvviiffoolliiuumm QQuuiissuummbb..&& MMeerrrr..
DESCRIPTIONS
4. Daphniphyllum ceramense(T.C.Huang) T.C.Huang, stat. nov.
Daphniphyllum glaucescensBlume subsp.ceramenseT.C.Huang,Taiwania 12(1966) 190,f.32.
—Type:Eyma 2047 (BO holo; K, Liso).
Leaves short, narrowly elliptic, apex cuspidate, baseacute-obtuse, 4-6 by 1.8-2.5
cm, coriaceous, shining onboth surfaces, lateral veins 6pairs,reticulationfine.Sta-
238 BLUMEA —Vol. 41, No. 1, 1996
minateinflorescences 2.5-4cm long, pedicles angulate, 5-7mm,calyx lobes4or
5, linear-ovate,4/5as longas androecium, caducous, articulate, stamens 8 or9,fila-
ments flat-oblong, 2-4 mm, anthers oblong to oblong-ovate, 1.2-1.6 mm long,
apiculate. Pistillateinflorescences 2-2.5cm long, calyx lanceolate,two whorls of
severalfreelobes, ovaryblue green,style branchesrecurved.Fruiting axes 2-3 cm
long, drupes smooth, ellipsoidal, green,laterblack purple, 1.3by 0.7 cm;fruitstalk
5 mmlong.
Distribution— Indonesia:Moluccas(Ceram).
Notes— Thespecies is similartoD. borneense Stapfexcept for having obscure
leafvenationand caducouscalyces onstaminateflowers. Thefree, lanceolate,two-
whorledcalyx lobesare verypeculiar.
ThenamereferstotheislandCeram
Specimens examined. MOLUCCAS. Ceram: first open slopeofMtMoerkele besar, Eyma2046,
2047 (L).
5. Daphniphyllum dichotomum(T.C.Huang) T.C.Huang, stat. nov.
DaphniphyllumglaucescensBlumesubsp. dichotomumT.C.Huang,Taiwania 12(1966) 190,f.31.
—Type: Clemens s.n. & Suppt. (US lecto;A,BO,L,NY, US, iso).
Daphniphyllumreticulatum H.Heine,FeddesRep.54 (1951)232,nonKeng(1951).
Shrub or small tree, 7.5 m high,60 cm indiam., bark smoothgrey, innerbark yel-
lowish, sapwood white yellowish. Leaves narrowly elliptic, apexacuminate, base
cuneate,7-10by 2.2-2.9 cm, coriaceous, greenonbothsurfaces, veins delicately
reticulateandprominent onboth surfaces, with4or5 pairs ofobscure lateral veins;
petioles 0.8-2 cm long. Pistillateinflorescences 2.5 cm long; calyx 4- or5-lobed,
lobes triangular, caducous, styles recurved; pedicels c. 2 mmlong. Fruiting axes
2.5-4.5 cm long, 1 mm thick; drupes ovoidal,7 mm long, 6mmthick withoutca-
lyx, greenandredbrownwhen ripe; fruit stalks 4-5mmlong, I mmthick.
Distribution—Indonesia:Borneo(Sabah, Sarawak).
Habitat& Ecology —Primary forestonlimestonecliffandboulders oforganic
soil atanaltitudeof960 m.
Notes—Thisspecies differsfromD.borneenseStapfmainlyby itsnarrowly ellip-
tic leavesanditsoccurrence at lowelevation. Itis closely alliedtoD. sumatraense.
Thenamerefers toits dichotomousbranchlets. Thevernacularnameis teradamu
bukit(Dyak dialect).
Specimens examined. BORNEO. Sabah: Kinabalu National Park,nearpark headquarters above
Helicopterlanding,Aban & PatrickLassan 79578 (L);Sarawak: GunongApi,Ulu Melinau,Tutoh,
Baram District, NEflank ofmountain,J.A.R.Anderson 30884(L).
7. Daphniphyllum gracile Gage
DaphniphyllumgracileGage, Nova Guinea 12(1917)480; T.C.Huang,Taiwania 12(1966) 155,
f.22;Barker inHenty,Handb.Fl. PapuaNew Guinea 2 (1981)34.—Type:Pulle 2439(Llecto;
BOiso).
T.C. Huang: Malesian Daphniphyllum 239
b. var. newirelandumT.C.Huang, var. nov. — Fig. 2
Avarietate gracilefoliis tenuitibus non-glaucisdiffert.—Typus:Gideon s.n.(CANB
462230,holo).
Fig. 2.DaphniphyllumgracileGage var.newirelandum T.C.Huang.A. Twigwith fruits;B.tuber-
culate drupe.
240 BLUMEA Vol. 41, No. 1, 1996
Shrub,c. 3m high, witharoundcrown.Leaves oblong-elliptic, apex obtuse,cuspi-
dateormucronate,baseobtuse, 6.5-10by 2.5-4cm, margins revolute, semi-coria-
ceous, lateral veins7pairs, membranous.Fruiting axes 2.5 cm long, calyx narrow,
caducous, styles short, divaricate, caducous; drupes green,ellipsoidal, 8-10 mm
long,7-8 mmindiam.,smoothto verrucose.
Distribution—Bismarck Archipelago, NewIreland,endemic.
Habitat&Ecology — This variety growson topofasmallhill(dominated by Glei-
chenia) whichis continuously burnt;altitude980 m.
Note—ThenamereferstotheislandNew Ireland.
Specimensexamined. BISMARCKARCHIPELAGO. New Ireland: Subprov.Konos,E ofLelet farm,
Gideon s.n. (CANB 462230,holotype);Lelet plateau,Gideon& Obedi s.n.(CANB 462228,para-
type).
11.Daphniphyllum papuanumHallierf.
Daphniphyllumpapuanum Hallier f.,Meded. Rijks Herb. Leiden 37 (1910) 13. —Type:Sayer
1887 (Lholo).
b. var. tuberculatum(T.C.Huang) T.C.Huang, comb. nov.
DaphniphyllumgracileGagevar.tuberculatum T.C.Huang,Taiwania 12 (1966) 159,f.22E; in
Henty,Handb. Fl. PapuaNew Guinea 2(1981)37,excl. specim. Sayer 1887.—Type:Brass
13705 (A holo;BO, L iso).
Shrubor small tree in understorey, 3-15m high, 2-24cmdiam.,outerbark greyor
brown, large pustular, brown green,scrape green,innerbark lightbrown, yellow,
blazeamberyellow, sapstainingwoodyellow, woodcream, whiteor straw-coloured,
yellow,buds green.Leavesoblanceolatetolanceolate, 5-20by 3-7.5 cm,apexacu-
minate,base cuneate,margins entire, hardchartaceous, lateralveins elevatedabove,
9-14pairs, not glaucous, innovationslight shining greenabove,adultleaves green,
glaucous ornotbeneath; petiole red, 1.3-5cm long,0.6-1 mmwide.Staminatein-
florescences4-5.5cmlong,calyxcaducous,stamens 8-10,antherselliptic, apiculate,
1 mmlong, 0.6 mm wide;pedicels 6-12mm long. Pistillateflowers with bluish
pruinose ovaries, stigmas bi-recurved, subsessile, purplish red, centre brownish
black. Young fruitgreen,shadedbrown or glossy light greenonapendant axis,of-
ten numerous; pedicels 6-8 mmlong. Fruiting axes 3-4.5 cm long,0.8mm wide,
drupes oblique ovoidtoovoid-ellipsoidal, greenthen darkbrown, black, tuberculate
orverrucose, 7-10mmlong,7-9mmwide;fruitstalks5-12mm long.
Distribution— EndemictoNewGuinea(Irian JayaandPapua NewGuinea).
Habitat&Ecology —In secondary forestonlimestone, burntarea, well-drained
volcanicsoil, in mixedprimary forest;broadridge forest;lowermontane rainforest,
ona Casfanopsi's-dominated ridge, andsubmontaneNothofagus rainforest; atalti-
tudesbetween900and2660m .
Notes— Thetype varietyofD. papuanumdiffersfromvariety tuberculatumby
thesurfaceofthe drupesonly; theformerhasasmoothto verrucosesurfaceandthe
latteratuberculatesurface.
This species is similartoD. glaucescens Blumebutdiffersonly by its caducous
calyx. Thenamereferstoitstuberculatedrupe.