The Species Diversity of the Genus Piper from Thailand

2008·7.1 MB·English

by  Chaveerach, Arunrat

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ISSN 1346-7S6S Acta Phytotax. Geobot. 59 (2) 1:05-163 C2008)
The SpeciesDiversity theGenus Pt'perfrom Thailand
of

ARUNRAT CHAVEERACHi, RUNGLAWAN SUDMOON2, TAWATCHAI TANEE3

PIYA MOKKAMULi
and
'Dapartment ofBiol oFgaclul;ty qf'Stienc eK,hon Kden Uhiversiij lKhon Kaen 40002, Tltaiiand '?Department of
Biochemisti yFaculty ofSZrie nIVcieo,n Kden Uhiversii jKlhon Kaen 40002, 7Viailan d'1'laculty ofEnvironment
StudiesM,dhasarakham U}iivensi4MeZihasarakham44000,77iaitand
andResouree
Species diversi toyf the genus Piper from Thailand was explored based on the herbarium specimens
kept at the BK and compared with the specimens deposite din the BCMU, BK, BKF, NX PE and
SING. [Ibxonomic keys available in the flor aofneighboring countries together with other taxonomic
literature swere used for botanical identificatio nF,rorn the investigation ,fort yspecies ofthis genus
were identifie dO.f these, two new species are described :R maculapIrylium and R rubroglandu-
losum. Four species: R betloide sR, clominantinervium, R pilobracteatum and R phuwuaense, in
addition with two varieties: R pedlcellatum var, agland"iatum and R thomsonii var, trichostigma
recently reported are included .The other thirty two species recognized are R argyritis, R bette ,R
boehmeriijbiiumR, R R hongkongense,R khasianum,R laetispicumR.lolot,
caninum, colubrinum,
R longum, R magnibaccum. R montium. R muliesua, R muricatum, R mutabile, R nigrum, R pecli-
ceUatum, R penduli,spicu mR, politij?)l iRum p,olyop{phonum R, retwfractum, R ribesioides, R sar-
mentosum, R semiimmersum, R submultinerve, R sylvaticu mR, sylvestn eA, thomsonii var. thomso-
nii, R tricoion R wallichii, R wangii. R yinkiangense; along with two unlmown species: Piper sp. 1
andPiper sp. 2 which may represent new taxa, will be furthe srtudied,
Keywords: New species, Piper, Piperaceae, Species diversity,Thailand
The genus Piper is estimated to consist of et aL 1963, Brown 1884a, b, Candolle 1910a, b,
over 1000 species, distribut emadinly in tropical Chaveerach 2003, Chaveerach & Picheansoonthon
regions of the world. Severa lspecies have great 2004, Chaveerach et aL 2002, 2006a, b, 2007, Y
economic and cultural importance includin gR C. Tseng et al, 1999, Chew 1972, 2003, Gard-
nigrum, R betle ,R methysticum, etc, Many spe- ner 2003, Gilber t& Xia 1999, Henderson 1959,
cies have been used for foods, medicines, stimu- Heyne 1987, Ho 1999, Hooker I885, Huber 1988,
lants, inThai Keng 1990,Lin & Lu 1996, Long 1984,Mer-
antiseptics and antioxidants culture
(Picheansoen ett haLo n2001). ril ]1912, Quisumbi n1g930, Ridley 1920, 1967,
Our taxonomic accounts on this genus are Roxburgh 1820) ,Royen 1982, fawan et at. 2002,
based on the stamen and stigma numbers and Tebbs 1982 and YUncker 1972. Moreover, there
characters, flora lbract morphology, leaf shape, are some reports taking molecular techniques into
and leaf venation (Y .C. Tseng et aL 1999, Jara- account to differenti aPitpeer species (Chaveerach
& Manos 2001).The identificatiofno1- 2002,Chaveerach2003).Recently,15Piper
millo et aL
lowed many references: Aghard 1981, Backer species were presente d(Chaveera &c hPichean-
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D MALAYSM
FIG. 1. The seven florist irecgions ofThailand: Northern (I )No,rth-eastern (II )Ea,stern (II IS)o,uth-weste (rInV) ,Central (V),
South-easter n(VI )and Peninsula (VII ()Srniti n&a nLadrsen 1972) ,and sites ofinvestigation < * ).
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soonthon 2004) ,Moreover, we have also studied deposite din the BCMU, BK, BKF, NY PE and
the ethnobotany of the genus Piper in Thailand SING have been studied. Usage of morphologi-
(Chaveera ceth aL 2006a). cal terms in this study is provided in Fig, 2, As
We have investigated the genus Piper for the result, currently, a total of40 species are
over ten years based on fiel dand hcrbarium stud- describe dand Mustrated. Of these, 32 species are
ies .Taxonomic identificati owenrse investigatedrecognized, fbur new species and two new varie-
based on references available, The investigationties recently reported (Chaveera ect hal. 2006b,
has covered almost all area of the country. Figure 2007) are included, In addition, two new species,
1 shows the seven fioristi cregions of Thailand R maculap]tyllum Chayeer, & Sudmoon and R
according to Smitinand & Larsen (1972 a)nd the rubroglandulosum Chaveer. & Mokkamul, are
sites of our investigatio nAs ,total of specimens firstl dyescribed in thi spublication
leaves onfree
branch

]rnalefiowers
bracts
1
/
/
a
g branch
FiG. 2. Usage of morphelogical terms.
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Key to species ofthe genus Piper in Thailand
IA. Spike straight up
2A, Small shrubs
3A. Fruitin gspikes 8-16 cm long d,rupe sarranged loosel y..,,.,,,,,......,,............ R.. s.y,l.v.e.s.t.n.e.,
3B.Fruitingspikes1-1.5cmlong,dmpesarrangeddensely.......,.........................,,,Asarmentesum

2B.Climbers

4A,Bractshairyadaxially............,.,.....,,....,,,,,,,........,,.,.,......,,,.....,..........Rpilobracteatum
4B. Bracts not hairy adaxially
5A.Plantsdioecious

6A. Leaf blade thin, ovate to lanceolate or narrowly evate; base often slightly unequal or cuneate
R caninum
6B. Leafblades membranous, ovate to circular; base cordate with rounded lobes ...... .R.. sylvaticum
5B. Plants monoecious
7A. Stigmas 34 on a spike ,.,....,,,,,,...,..,,,,.....,,,.,,,,......,,,.,,.....,, ,m,ul,te,s.ua....,,.........R
7B. Stigmas 3 on a spike
8A. Fmiting spikes O.72.5 cm long ;drupes globose ,green when ripen .........,,...... .R, .J.o.n.g.um
8B. Fruitin gspikes 3,5-6 cm long ;dmpes globose, red when ripcn , . . . . . . . , , . . . . . . . . . . . . . . R retrct17"acturn
1B. Spike pendulous
9A. Leaves on epiphytic branches ,adaxially dark pink dapple ,abaxially green . . . , , , , , . , , . R maculapdyllum
9B. Leaves on epiphytic and free branches both sides or a side pinkish-lig ghrteen ,ligh tgreen or gTeen
1OA, Plant sall part shairy
11A. Small shrubs ...................,.......,.......,.....................,,.... .m.ur.ic,a.tu.m...,,.....,,,,,...R
11B. Climbers
12A,Leafb]adesovate;basecordatewithunequallobes......,,..................,.........Ryinkiangense
12B. Leaf blade sovate-lanceolate, oyate-oblong, obleng; base oblique, rounded or subcordate with
lobes divergen tor overlap each other
13A,Stamens3.......,.,,...,,,,,,......,,,.......,,.,....,.,,,,........,,,,.,,..,,........,...Rhongkongense

13B,Stamens2

14A. Leafblades ovate-lanceolate, ovate-oblong; base oblique, rounded or subcordate with lobes
diverge nort overlap each other; veins 7, two pairs basa l....,.... R.. s.emiimmep:sum
14B. Leaf blades oblong; base deeply cordate to auriculate or overlapping auriculate; veins 9, three
pairsbasal,.............................................................................,..Rsubmultinerve
1OB. Plants glabrou sor some parts hairy
15A. Fruit swith stipe
16A.Stamens4,...,......................,,..........................................................Rribesioides

16B.Stamens2..,.,,.......,.......................................,...........,,......,,.......,,...Rpolitijiolium
15B. Fruits sessi]e or stipe not seen
17A. Bract swith censpicuous stalk, protruding stick out ofrachis ................. .R. .p.e.d.i.cellatum
1 7B. Bract without stalk or without conspicuous stalk
18A. Small shrubs ,,.........................................,,........ .R. .b.o.e.h.m.e.ri.i.j.2.].l.i.u.m.....
18B.Climbers

19A. Leaves finel ypubcscent or densely glands
20A. Leaves abaxially finel pyubescen talong veins, veins 5, all basal ................... .R. .l.o,l.o,t.
20B, Densely brownish-re dglands along veins, veins 7, two pairs basal a, pair arising 2-4.5cm
apartfrombase,oppesiteoraltemate,,.......,,,.,....,,,......,,,...,..,,.......,......,.Rthomsonii
20C. Leaves ahaxially and adaxially densel yscattered white glands . , . , , , . . . . . . , . , . . . , . . . . . R betloides
20D. Leaves adaxially densely scattered reddish glands ...,,,,,.,,....,,., ,R. .r.ub.ro,g,ta.ndulosum
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19B, Leaves abaxially not as above
21A, Bracts divide dinto areas; center black ,surrounded by yellow margin ........... .R. .t.ricoior
21B. Bract snot divide dinto areas
22A. Bract base sadnate to rachis; stamens 3, male fiower ssparsely scattered on rachis, filament
fiat l,onger than anther, raising the anther up above bract ....,............ .R. .p.o.l.y.syphonum
22B. Bract bases not adnate to rachis
23A, Ovary arising from hole of rachis .,,,,...........,,...,,,,.,,........ .R. ,k.h,a,s,i,a,n.u..m...
23B. Ovary not arising from hele of rachis
24A. Bract margin ciliate
25A,Stamens2..........,.,,,...................................................,..Rclominantinervium

25B.Stamens3

26A. Stem node stout and dilate .d....................,.,................ .c.ol.ub.r.in.um...,,...R
26B.Stem dilated
neither node stout nor
27A. Male spike 7-1O cm long ............,.....,,,,,.,............... ,l,a,e,t.i.s.p.i..c.u.m..R
27B, Male spike 2-3 cm long ..,,,,,,,,,,,,,...........,,,,,.,,.......,,..,,,,.,,,,Rphuwuaense
24B. Bract margin not ciliate
28A. Plant smonoecious
29A. Male and female fiower son a spike .................................. .P. .n.ig.r.um........
29B. Male and female fiower son separate spikes
30A,Stamens3....,,,.........,........,..,,,.,,.........,,,,,,,,,,,..........,,,,,,,....Rwallichii
30B.Stamens2..............................,,,,,...........,..,,,,...,.........,,.Amagnibaccum

28B. Plants dioecious
31A,Leafbladeswithpaleorwhitespots.....,..,..,,,,,,,....,...,..,.,,,,,............Ra,gyritis

31B. Leafblades without spots
32A.Veins5-7

33A. Leaf blade selliptic ...........,...........,..,,,..,,........,,,. .R, ,mo,n,ti.um.,,....,....
33B. Leaf blades narrowly ovate to evate .......,,,..............,.,.,. .R. .m.ut.a.b.il.e...,
32B.Veins6-7,7 7-9
or
34A. Stigmas 4-6 ..........................,.,,,,,...........,,,,,..... .R. .b.e.t.l.e....,.........
34B.Stigmas3--4

35A. Female spikes 1O-20 cm long .............................. .Rp.e.n.d.u.l.i.s.p.ic.u.m
35B. Female spikes 1-2 cm long ..........,,,,,,,..............,,,,,,. ,R.. .wa.ng.i.i.........
Enumeration Species
of
1. Piper argyritis Ridl, ex C. DC., Rec. Bot. spike pendu]ous, 6-8 cm long, O.2-O.3 cm in
Surv ,Ind ,6: 25 (1912 ()F,i g3). diameter ;peduncle 1,2-1. 5cm long ;brac torbjcu-
lar ,abaxially pubescent ,adaxially convexed; fila-
Dioecious ,stout climber, node slightly dilated.
ment flat ;starnens 2. Flowering on year round.
Pceeotusi,ol eg l1a-b1r.o5u scm, l ilgohn ttgo; dlaerafk bglraedeen , tphailcek orch awrhtiat-e Local name. Sa-kan, Sa-kan-hin ,Sa-kan-
dook.
dried, 4,5
spots easily seen when oblong-ovate,
-5.5 12-15 long; Distribution .Northern Thailand, 800-900 m
cm wide, cm apex acuminate;
alt
base oblique, slightly cuneate to rounded; veins
elevating beneath ,one or two pairs basal ,the oth- SPeeimens examined. THAILAND: Northern,
ers arising from midrib at regular interva lM.ale Chiang Mai province ,cultivated in viIIages, 900 m alt,,
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October 12, 1998, A. Chaveerach 11 (BK 63490); Bei- slender or stout; node of yeung branch inflated,
1'ingyoung group 1467 (PE 680455). O.5-O,7 cm thick, Petiole not sheathed at base,
Notes. This species is cultivated in villages 1.5-2 cm long. Leaf blade leather ythick, with
fbr woody stem used in soup, but not as popular white glands densely scattered on both surfaces,
as P pendulispicum due to it sstem is harder. dark green ,narrowly ovate to ovate, elliptic or
5.5-17 11.5-25 long;
obleng, cm wide, cm apex
2.Piper betleL,,Sp.Pl.1:28 (1753)(.Fig4.) acuminate; base obliquely truncate or obliquely

rounded; margin entire; veins abaxially promi-
Dieecious, Stem
climber. stout with pinkish-stripe nent, twe pair sfrom the base, fou ror five pairs
along, node dilate dand rooting. Petiel e2-2.5 arising from midrib. Male spike unknown, Female
long;leafbladefleshy
cm coriaceous, glabrous, spike pendulous, white, 2.5-3 cm long, O.5-O,7
greenish or yellowish, broadly ovate, 7-8,5 cm cm in diameter ;peduncle 1.5-2 cm long ;bract
9-11 long; base
wide, cm apex acuminate; cor- elliptic, shortly stalked, base pubescent ;stigma 3
date ;veins 7-9, elevating beneath ,two or three -4, hairy Drupe globose ,loosel yinterrupt eodn
pairs basal ,one pair arising from midrib. Male rachis. Flowering and ffuitin gon May to June.
3-l2 long,
spike cylindric, slender, pendulous, cm Local Plu-doi-su-thep.
name.
O,5 indiameter; 2-3 long;
ca. cm peduncle cm Distribution,NorthernThailand, Hill
ever-
bract orbicular, peltat es;tamens 2. Female spike green fores ntear water fa11 1,1OO m alt,
cylindric, pendulous, 2,5-4 cm long ,ca. O.5 cm in
diameter ;peduncle 2-3 cm long ;bract orbicular, SPecim eenxamined THAILAND: Northern, Chi-
p5 eclmta lto ens;tgi;g mdasru p4e -e6m,b epdduedbesc eonnt r.Farcuhiist.j nFg lsopiwkeer i3n-g am6n3g 7a2 l0Mt)a.,, iM apyro v1i5n, c2e 0,D0o1i, AS.u tChheapv-ePeur iaNcaht i4o7n a( lBPaKr k6, 31511O4O,
and fuiting on year round,
Local name. Plu. 4. Piper boehmeriifolium (Miq. W)all, ex C. DC.
Distributi oWnh.ole Thailan dto China, Indo- in A, DC., Prodr. 16(1): 348 (1868 )H;ook f, in
nesia, Malaysia ,Philippine sSr,i Lanka, Vietnam, Fl, Brit. Ind. 5: 85 (1885 )R;idl, ,Fl. Malay Pen.
Madagascar, 3:41(1967)Ho;,IllustF,l,Viet,1:293(1999)-.

Chavicaboehmeriij?)iMiiqa., SystPiperac.:265
EPecime enxsamined THAILAND: North-eastern,
(1843)(,Fig5.)
Khon Kaen province ,200 m alt., October 22, 1998,
A. Chaveerach 16 (BK63494); ei-wuPTZing75416
(P E200005554) ;H / IVL RidtqySf fVG468 4 (SI NG)Z; Dioecious, slender shrub, up to 2 m tall. Stem

7lem{ySa7NG 869 (SING). warted, internod eup to 30 cm long ,young stem

hairy.PetioleO,5-1 long; leafblade
cm mem-
?Vbtes .This species is cultivatgd for tradi- branous, lanceolate ,4-7 cm wide, 17-23 cm
tiona lmedicine. Many Thai people are familiar long;base
oblique with unequally auricles; apex
with this plant because it is always seen in several acuminate; veins 7-9, one or two pair sbasal ,the
traditional activities. Its leaf is used fbr wrapping others arising from midrib, alternate, reticulate
ofbetel nut and lirn efor mastjcatory. veins transverse ,Male spike 10-16 cm long ,O.2-
O.3 cm in diameter p;eduncle 1-3.5 cm long ;bract
3.PiperbetloidesChaveer.& Tanomtong, Acta
orbicular, inserte dbasall yor peltate ;stamens 2,
PhytotaxG.eobot,58(1):33-35,fig,1(2007).
Female spike 6-12 cm long, ca. O.2 cm in diam-
Dioecieus, climber, fragrant like R betle .Stem eter; peduncle and bract as in rnale spike; stigmas
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C
B
F]G. 3. Piper at gyriti Rsidl .ex C. DC, A: Branch with male spike. B: Part ofmale spike,C: Male fiower.A. Chaveerach 11
(BK63490).

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';
"f''.J-
ts
u
I2cm
L
- p- --
Iicm
B
A
FJG .4.Piper betl eL, A: Free branchwith fernal espike.B] Part of female spike.A. Chaveerach 16(BK63494).
deciduous .Fruitin gspike 6-12 cm long; drupe IVimanong & S. Phusomsaeng SllVG 269 (SING) Z;E
subglobose, arranged densely on rachis. Flower- Mbxwell BKF 046689 (BKF) ;Pino,i ltdi n3o738 (PE),
ing and fruiti nfgrom November to March. 9i-xi Lniu 26384 (PE 21OOO1662); Mlliam BKI' 17133
(BKF),
Distribution ,Whole Thailand te Bangladesh,
India,Myanmar, China,Bhutan,Malaysia,Sik-
kim,V ietnam,10 0-900 5. Piper caninum Blume, Batav. Gen. 11: 214
m alt. (1826()F.ig6.)

Specimens THAILAND: North-
ea st er n, Phu Luan gex aWmiinledd.lif eS anctuary ,Loei p rovince, Dioecious, slender climber. Stem fragile ,gla-
900 m alt., July 9, 1998, A, Chaveerach 2 (BK 63480), brous, sparsely short hispid .Petiole slender, frag-
September 2, 2004, A, Chaveerach S5 (BK 63480); B. ile ,O.5-1 cm long ;leafblad ethin, membranous,
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base; 5, basal,
or veins one pair another pair aris-
ingfrom base.Male
midrib near spike straight
up, 1,5-3 cm long; peduncle slender, fragile ,O.5
-1 cm long ;rachis pubescent ;brac tsuborbicular,
with not smooth edge; stamens 2, with long fila-
ment, exerted from bracts ,Flowering from April
toOctober.

Distribution. Southern Thailand to India,
Malaysia, 100-600 m alt.
SPecime nesxamined. THAILAND: Peninsula,
KhaQ Maen, Nam Tok Ybng National Park,Nakhon

Sri Thammarat provinc e1,00 m alt. ,January 2, 2004,
A. Chaveerach79 (BK);A.EG. Kerr 11747,i2567,

J3312, 13663, 16732,18347, 18561 (BK);E. J.H.

SJNG 28492 (SING)H;. R. Henderson SI7VG 12822

(SING)Z;S.BurlqySfiVG3776 (SING);T Sinittinand

BKF 100f (BKF).

6. Piper colubrinum Link, Jahrb. 1(3) :61 (1820).
(Fig7,)

Dioecious ,scandent, 3-4 m tatl .Stem stout joint,
node dilate adnd rooting, 2-3 cm in diameter P.et-
iol eO.5-1 cm long ;leaf blade coriaceous, papery,
glabrous l,igh tshiny green ,ovate to lanceo]at 7e.,0
-8.5 cm wide, 17-18 cm long; apex acute; base
9, basal,
oblique; veins one pair one pair arising
O.3-O.5 cm apart from base, the others arising
altemately from midnb with ene pair reaching to
apex, Male spike pendulous ,white cylindric, 4-
5 cm long, O.6-O,7 cm in diameter; peduncle ca. 1
cm long; bract elliptic with edge hairy; stamens 3;
Flowering on year round.
Locatname, Prik-thai-tua-pu,

Distribution.Whole Thailand,

FTG .5 ,Piper boehmeriijlo l(iMuimq. C). DC. fipecimen sexamined THAILAND: North-eastern,
fruitin sgpikes, A. Chaveerach 2 (BK 63480).Branchwith Khon Kaen province, 200 rn alt., October 22, 1998,
A. C)haveerach19 (BK);X E Maxwell BCMU 96-858

(BCMU);T K.Kueh,Si?VG153(SING),
variable in shapes and size, ovate to lanceola toer
narrowly lanceolate ,rather small, 2-4.5 cm wide, ?Vbtes .This species is cultivated in lowland
7-8 cm long ;apex acuminate; base often slightly garden usually used as a stalk fbr grafting of P
unequal or cuneate; veins 3, one pair arising near nlgrum.
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TThhee JJaapapnaesneSeosciee tySociety ffoorr PPllanatnt SSyystsetmaetmicastics
114 Acta Phytotax .G e o bo t . M)L 59
7. Piper dominantineryium Chaveer. & Mokka- oblique with unequally sides, auriculate. Veins
mul, Acta Phytotax. Sin .44(4): 447--449 ,fig .1 differe nftrom most Piper species, many, all aris-
(2006), ing from midrib at regular interval osf base to
joining
Ddiiosetciinocutslj ,yocliimbnert, on trbereos w.Sntiesmh -sbtoluat,ck niosdehs aoptehxer. r eMacahlineg sp inkeear pmeanrgdiunlou asn,d 5 -13 cm w iltohn ge, accah,
-feob9sul orpnregtoiw loetolod e w n hng1eanr;-r1ow .l dy2 srcwimo lele de llnL,l,oeinpagtv b;ie acsl ,se 1eoa.nf5 -be3pli.ap5h dycet micco rwii adbceer,oau s na,7ncd.h5- ObFto.er 3maN acoclmt veeel mlsiibppnit eikdrcei,. a nmosteest sseierlee;,n .p emFadlrgouiwnnecrli en ciOgl. if7atr-eo1;m. 5 s Stacemmepnt sleomn b2ge.;r
cm apex acute; subcuneate-cuneate Locai Plu-tum-nang,
or subrounded. Leaves on free branches petiole Distri nbaumte.ion .Souther nThailand.
O.1-O.2 long;leafblade dark
cm coriaceous,
green, densely yellowish, brownish or reddish thecime enxamined THAILAND: Seuthern ,Sri
glands ,ovate or narrowly elliptic to elliptic, 2.5- Phang Nga Nationa lPark, Phang Nga province ,100 m
9 cm wide, 12-19 cm long; apex cuspidate; base alt., October 19, 2000, A. Chaveerach 43 (BK 63498,
63499,63500),

I2cm
FJG. 6. Piper caninum Blurne.Free branch with fiowering spike. A, Chaveerac h79 (BK).
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