Variation of Pitcher Morphology within Nepenthes vieillardii Hook. f. (Nepenthaceae) in New Caledonia

2004·2.1 MB·English

by  Kurata, Kaoruko

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ISSN1346-7565 ActaPhytotax.Geobot,55(3):181-197(2004)

Variation PitcherMorphology Nepenthes
of within vieiUardii

Hook. fi(NepenthaceianeNe)w Caledonia

KAORUKO KUR-ATAi JAFFRE2 HIROAKISETOGUCHIi
and
,TANGUY
'Graduate SZrhoo olfHLtm aandn Environmental Studies ,K)Jo tUhoive,isiC B1<l,oto606-8501 JZ{pan; ? Researeh
lnstitutefo rDevetopment, BP A5, IVbutnea, ?Vlew Cdtedonia, Fhance,
[Ihxonomy of Nbpenthes has been exc]usively based on pitche mrorphology that well characterizes the
carnivore. In order to understand the morphological divers iantdy to rc-cxamine taxonomical diagno-
sis ofthe Nepenthe staxa in New Caledonia t,he pitche mrorphology was carcfUlly studicd based on 221
individuaflrsem localities islandF.orthis inves-
matured upper pitcher soM24 seven ofthe purpose, we
tigated size ofpitchcrs and oftheir lids n,umber and densit yofnectar glands on lids n,umber and den-
sity of digesti gvleand sin pitcher asnd ratio of digestiv zconc vs. waxy zene in pitcher sT.hese mor-
phologica fleatur evsaried widely betwee nthe populatio ntshough they are mostly stable within an indi-
vidual plant .Statisti canaallyses did not segregate any populatio annd supported the taxonomical treat-
ment to unite several taxa int oan insula erndemic, 7Vlepenthe vsieillaJxlii Hook. f in New Caledonia. The
number ofdigestive gland sand area ofdigestive zone in pitcher varied greatly between populations sug-
gestin gthat some ecolegical factor dso affect the morphological diversi toyfpitchers.
Key words: carnivorous plant sd,igesti vgelands m,orphology, nectar glands ?,Vepenthe Nse,w Caledonia,
pitcher
Nepenthes t,he sole genus ofthe famil yNepentha- shows a wide range in it smorphological diversity
ceae comprises about 90 species ofperennial herbs (e.g M.a,cfarlane 1908, Kurata 1976, Komiya 1994,
distribute din Madagascar, Seychelles ,northern Phillip s& Lamb 1996, Clarke 1997, Jebb & Cheek
India ,Indo-China peninsula ,Malesia ,northern 1997) .Therefbre t,axonomical confusions have
Austral ianad New Caledoni a(Kura 2t0ae2) ,These been caused in the current classification of
plant scould expand thei rdistribut biyo nadjusting to AJkipenthes.
soils with poor nutrients owing te an accomplished New Ca]edonia, a continental islan dthat orig-
insectbsy insec- inatedfromGondwana,is latge
special evolution ofcapturing an covered with a ser-
tivorou sleaf called a "pitcher". Taxa ofNepenthes pentin earea, The ultramafic soil derive dfrom the
have been mainly classified based on the charac- serpentine or gabbro is well characterized by very
teristi cofspitcher morphology, On the other hand, few nutrients fbr plant growth and hypeFmagnesium
dimorphism of the upper funne lpitcher and the content and the IVepe nthes plant is well adapted to
lower janin pgitche rhas been well recognized even the edaphic conditions. Concernin gthe species in
within an individual p]ant (e.g .Cl,arke 1997). New Caledonia n,ine taxa including six endemic
Intraspeci fvairciation of pitche mrorphoLogy also species have been describe dmainly based on the
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morphology of the pitcher (Hooke r1873, Dubard tine area with poor nutrient. The populatio nexsam-
1906 ,Macfarlane 1908, Guillaumin 1911, Moore ined in the presen tstucly were chosen ftom near]y
I921, Danser 1928, Guillaumin 1953, 1964, Jebb & entire area of distributi roannge of the species,
Cheek 1997). These taxa of Nepenthes in New moreover, fiv eofthe seven population esxamined
Caledoni haave been describ ebdased on small num- were locate din the southern part ofthe islan d(Fig.
ber of specimens, and i tis uncertain whereabouts in 1 and Table 1-A) where the serpentine soil is wide-
the plan tbody the pitcher wsere collected. ly distribute (dORSTOM 1983), Population ofGR
Moreover, Jebb & Cheek ( ] 997) integrat ealdl is devclope don gabbro and the remainders on ser-
taxa reported from New Caledoni aas, a synonym of pentmc.
?Vbpenth evsieillardii Hook, fl without any expla- Prior to the analysis between the populations,
nations. Further examination ofthe morphological morphological variation ofpitcher within an indi-
diversit aymong New Caledonian Nqpenthes is vidual was examined. Plant sof Albpenthes vieil-
needed to clarify the morpholegical diversi tquyan- laJdi iusually have several matured pitchers and
titatively and to verify the classification of the we could collect many pitchers ,enough fbr statis-
specles. tica lanalysis at CR GR, MK among the seyen pop-
As contribution to a critical cxamination of ulations and additionai RA and BL CTab l1e-B).
the taxonomic prob]ems of IVepenthes in New 14 - 28 pitcher wsere sampled from each individual
Caledoni aw,e clarified in thi sstudy the variation of in the fiv ecollection sites.
pitche rmorphology in Nepenthes vieillaFdii sensu Fer analysis on the inter-populati voarniaatlion
latoinNew Caledoniaby
quantitativ aenalyses on of pitcher morphology, we collected a matured
the size of the pitche rand it slid ,the number and upper pitche rfrom each individual ,and eight to
densit yof the ncctar glands and digesti vgelands, 28 matured pitcher wsere sampled from each local-
and the ratio ofthe digesti vzoene and waxy zone in ity .In total, 119 pitcher swere used fbr the mea-
the pitcher .Classificati oofn Ncw Caledonian surements ([la b1l-Ae). Since remova] ofpitchers
IVbpenthe ghas been discuss ebdy comparison among was expected to seriously damage thc individual,
the collection sites ,The nomenclature tentatively pitcher swere collectcd from large individual swith
fo11ow sthat ofJebb & Cheek (1997). many pitcher sT,he collected pitcher wsere fixed
with FAA (50% ethanol: acetic acid: fbrmali n=
Materials and Methods 90: 5:5).
S t u d"vPla ntssites and materials iWeasurements
ef Nepenthes vieillardii grow in maquis veg- Seven morphological characters were measured
etation (Broo k1s987) develope don ultrabasic soil (Fi g2.)i sizc of lid (maximum and minimum diam-
o.riginating from serpentine rock or gabbro. The eter of lid )l,engt hand width ofpitcher and height of
plant susually develo psmall populations isolated digestiv egland zone were measured using a scale;
from each other. NOpenthes in New Ca]edonia usu- the number and densit yof nectar glands on the lid
ally grown in sunny place lik ethat ofopened slepe, and digesti vgeland sin the pitche rwere ¢eunted
riverbank or edge ofthe fbres tT.he plant is creep- under a stereoscopic microscope (Olympu SsZ4045,
er ef several meters in length a,nd develop sleafi Olympus Co., Ltd.) .We did not use characteristics
pitcher son the top ofshoot while withering lower such as color, shape and presence or absence of
leafpitche rsTh.erefbre the plant sfi'equent lhyave the fring efbr this analysis because the pitchers
several matured upper funne lpitcher sin sexpen- showed larg evariations even within onc plan t(Fig,
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"ts

'O

KO

COL DE PRONY GORO(GR) ILE DES PINS
(cp) (IP)
FiG. 1. Map showing the location osfnine collection sitcs of Aiepenthes vieiUardii in New Caledonia.
TABLE 1. Col]ectio nsitcs and samples ofNepenthes in New Caledonia,
A. Inter-populationak,aria otfipiotncher morphology.
Code forst u dy ' Altitud e(m)Soil typeNumber of
sitcLocalitv pitcher esxaminedHabitat
IPGRCPMKIRlBeKdNeKsPPins 30 Serpentine IO28152821W9e8triverbank

GoroCol 803204202G2a0b8b3r0o81S0erpentine Open slope
d e P r on v' Open slope
Mont Koghi Serpentine ShtvbEdge

RiviereBleue Scrpentine ofthc forest
Kuniambu Serpentine Open slope
Kopeto Serpentine Edge ofthe forest
B, Variati oofnpitcher morphoLogy within onc plant.
Sample No,Loca[ity Code fbrstudv typeNumbcr of
site pitchers examinedHabitat
'Soil
12345 Mont Koghi MKBLGRcpRASerpentine 2814152520 ShnibOpen

R{viere Blanche Serpcntinc slope
GoroCol GabbroSerpentine Open slope
de Prony Opcn slope
Radier Serpentine Open slape
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A

H
<S.
e e
B
.
e ee
c
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<l e iliS)
e
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paee

FiG ,2, Measuremcnt ofpitchers and lid sof Akzpenthe vsieittardl'i A, Length ofmaximum djannet eorf Lid B., Length ofminimurn diam-
eter of lid C,. Ik)tu nlumbcr of nectar glands .D. Length ofpitcher. E. Width at the widest part ofpitcher. F, Height efdigestive
gland zone ofpitcher. G.. Number of digesti vgclands per unit area.
3). Thc area of the lid swas calculated from the Guillaumin includin gtheir type specimens were
rnaximum and minimum diarnete arnd, the density examined fbr measurcment of their pitche rmor-
ofthe ncctar gland swas estimated. The densit oyf pholegy. The available specimens were the four of
digesti vgelands pcr square was calculated from the nine taxa describe dfrom New Caledonia, 26
the area of the leafpiec ceut-off by the leafpunch. pitcher sfrom 12 sheets ofN vieillanlii, 15 pitchers
Specimcns ofAJepenthes vieillantii, N, vieil- from six sheets of N vieillardii var. dqplanchei,
laFdi ivar. deplanche iDubard, Nl vieillardii var. fiv epitche rfsrom one sheet ofAL vieiliaptiii var. min-
montrottzierii Dubard and N. vieillantii var. minima ima and 13 pitcher fsrom three sheets of ?V lvieil-
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ee
':-rtrv,,ll･Il
iffme
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･･,/,/,/tvt'/･t;/1,
//mmbt'
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FiG .3, Gross rnorphoiogy ofpitchcrs ofIL'(7]enthe,s vieillantii Thc color and the shape erthe pitche rvar-
ied ameng p!ant sT,he arrows indicat tehe boundaty between thc cligcstiye gland zone and waxy zone.
a, b and d were from RB, and e was fror nMK. f: indicate funge.
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tardi ivar. montrouzierii were examined for the given in Table 2-B. Length and width ofpitcher on
rnorphology. List ef specimens examined were list-plants from KP are large rthan these from other
ed in appendix, localiti e(smea n13.5 cm in lengt hand 4.4 cm in
width in KP). On the other hand, pitche irn plants
Statisticalanalysis from GR., MK and KN were relatively smal]er;
Statisti acnaallyses were performe dusing the mean width ofthe GR specimens (2, c3m) was almost half
value for the characters of each individua lT.he of specimens from KR
data were subjected to analysis ofvariance (one-way
ANOVA) with the Tukey' shonestl ysignificant dif 2) Nectar gtancls
ference test (Tukey H'SsD) using SPSS ver. 1O,O J Number of nectar gland son the lid of plants from
fbr Windows (SPS S2000). KP was markedly higher than those from other
localiti(e1s45.1 lid)l;essthan
glands on one-third of
Results
gland swere distribut eond the lid of specimens
fromGR (Tabl2e-B),

PZiriati oonfpitch meorrphotogy within one indi- Thc area of the li dwas calculated from the
vidual and between individuats lengths of the maximum and minimum diameters,
The seven morphological characters were measured and the densit yof the nectar gland on the lid at
for 1 02 pitcher sfrom fiv eindividua lansd, the mean each cellection site was also estimated. Intrapop-
value and standard error of the measurement of ulational variation and comparison among the pop-
each plant were presente idn 1labl e2-A, The stan- ulatiens in the densit yof the nectar gland on the 1id
dard errors for each measurement in individual are presente din Fig. 6. The value showed a varia-
were lower than the value ameng the individual tion range of 17.4 (KN) - 7,7 n,/cm2 (RB) i,ndicat-
(Fi g4.), The mean va]ue is remarkably differenting grea tvariability,
from each jndividu aNle.ctar gland densit yranged The relation between the area of the lid and the
from 6,6 (sampl 4e) to 24.3 nos,lcm2 (samp l1)e and number ofnectar glands is given in Fig, 7, A cor-
was remarkably high in sample 1 among the five relation was recognized between these two factors
individua lsO.n the other hand, ratio ofdigestive (Correla tcoiefofincient - O.5271 ,p < O.OO1) .The
gland zone f length ofpitcher was O.59 in sample 2, number ofnectar gland swas changed in propouion
indicatin gthe highes tvalue among the fiv eindi- to the area ofthe lid .However, the densit yofthe
viduals exarnined (Fi g5),, The significant difibrencenectar gland sis an unstable character on account of
was detecte bdetween all individu pallant sexcluding the iarg evariatjon within a population.
the densit oyfthe digestiv egland (p < O,O 1, Tukey's The distribut iofon nectar glands is shown in
HSD, Table 3-B), Fig .8-a .The nectar gland sof?Vlepenthes vieillardii
were sparsely distribute tdhroughout the entire lid.
lhteripopulationalvariation
D Size ofti adndpitcher sy Digestive glancts'
The variation ofthe area oflids is shown in [Ribl e2- The densit oyfthe digesti vgelands t,he tota 1number
B, The area oflids in KP was significantly ditTerentof digesti vgelands and the ratio of the digestive
from that in other population sT.he mean value is gland zone are also shown in [labl 2e-B,
large r(mea n14.4 cm2) than twice ofthose in GR, The yariation range in the densit yof digcs-
CP and MK (mea n5,6 - 6.6 cm2). tive gland son a pitche rwas 442.0 g]and sper cm2 in
Variation sin the size of the pit¢her are also IP to 733,7 glands in GR (Fi g9,) .No significant dif
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December　2004 KURATA 　et　al，；Variatio　nofpitcher 　morphology 　withinAN 　vieilla，dii 87
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188 APG Xio15.5

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xutivv30252015105
[os O,5
a
Ltu
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o
M,e
c[ooa 1(MK) (B2L3 ) (GR ) (4CP) 5(RA) ( 1MK ) (B2L ) (3GR ) (C4P ) (R5A)
(SCamopllle enucmbt esriitoen) S(aCmopllle enucmbterionsite)
Fi( }4.. Mcan and standard crror ofdensity ofnectar glands col- FiG. 5. Mcan and standard error of ratio ofdigestive gland zone
lecte dfrom fiy eindividual plants. co]]ectcd from fiy eindividua pllants.
TABLE 3.Significa ndtifferenc ocfmorphological characters in one plant using Tukey' HsSD test, (*p<O, 0**5p<,O.O1)
A.Density
ofncctar gland
Sample No. 1 2 3 4 5
12345
o.ooo**o.ooo**o.ooo**o.ooo**
O.11IO.O16*O.945
o.ooo**O.O08rk*
O.064
B.Density
ofdigestive gland
SampleNo. 1 2 3 4 5
.]2345
O.054o.ooo**O.355O.87S
O.39SO,762O.389
O,O15*O.O03*s
O.942
C, Total number of digestiv egland
Sample No. 1 2 3 4 5
12345
O,O06**o,ooo**o.ooo-*o.ooo**
O.Oooe*o.ooo**o.ooo**O.ooos*o.ooo**O.041*
D. Ratie ofdigestive zone
SarnpleNo, 1 2 3 4 5
12345
o.ooo**O.074O.O12*O.435
o,ooo**o.ooo**o,ooo**1.000O,O02**o.ooo**
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and the maximum width of a pitcher (Correlation
ouAE 25
Ux coeMcient =- O.7446 ,p < O.OOI). Therefbre we
dNL2Y0v used lengt hof pitche rfbr representing the size of
cre 15 pitcher in furthe srtudy. On the other hand, corre-
a latio nwas not recognized between the size of the
L
PaUs 10 pitcher and the density of digestiv eglands
oc-p 5 (Correla tcoieorancient - -O.214, p < O,05 ,Fig.
o 11).
M't
c[ooa IP GR CP MK RB KN KP The shape of the piteher was assumed to be a
Collectio snite column for estimating the total number ofdigestive
gland sin a pitche rbased on the maximum irv'idth of
F i G ,6 , M ea n a nd s ta nd ar d e rr or of d en s itvt of nectar g-lands col-
lecte adt seven localities. the pitche rt,he heigh tof the digesti vgeland zone
and the densit yofthe digesti vgeland ,The total
number ofdigestive gland son the pitche orfplants
from KP was significantly highe rthan those from
250Adve
other localiti e(smea n40,231. 1glands on pitcher);
2oog-en more than twice ofthose at IR GR, CR MK and KN
(mea n16,398. 6- 22,861. 7glandslcm 2a)s shown in
150ds82 Table 2-B. The variation and comparison amQng
populatio nisn the nurnber of digesti vgeland son the
pitche arre presente idn Fig .12 .The tota lnumber of
loovotsD
digestiv egland sshowed a relatively strong corre-
50E=z latio nwith the area ofthe digesti zvoene (correlation

coeMcient - O,7763 ,p < O.OOI ,Fig, l3),
oo On the other hand, no significant difference

10 20 30 was observed in the ratio of the digestive gland
zone (digest ziovneeftotal length ofpitcher) among
Area of lid s(cm2) the collection sites except at CP and MK (Fi g1.4,
[labl e4-D), The maximum mean value was O.47 at
FIG. 7. Diagram showing corrclation between the area of' lid sand
the number ofnectar glands ofthe pitche rat cach locality.IP, and the minimum was O.37 at CP. The mean
Correlatio nindex = O.527 1, value of the ratio of the digesti vgleand zene was O.5
or les s{n all the sites, i twas clarified as character-
istic osf Aiepenthes vieiUaidii that the digestive
fercnce was recognizedi in the densit yofthe diges- gland zone was distributc bdclow the lower halfof
tive gland sdetecte adt each collection site, howev- the lengt hofthe pitche rH,owever, thc distribution
er, significant difference wsere fbund between in JP (dept ho)f the digesti vgeland zone varied among
and GR, GR and MK (lbke HyS'D, p < O.O1, Tbble each pitchcr (Fig 8.-b).
4-B) .To examine the relation between the size of
the pitche rand the densit yof the digestiv geland ,the Measttrement oj'Hlerba srpieucimmen
correlation between lengt hand the maximum width 59 pitcher sfrom fbur taxa of New Caledonian
of the pitcher was examined (Fi g1,O), A relatively Nbpenthe swere exarnined based on thc hcrbarium
strong correlation was recognized between length specimens includin tgheirtypes ,Averagc and stan-
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TThheJea paJnaespeSaoncieestyefo rSPloacntiSyestteym atficosr Plant Systematics
190 APG 5,bl, 55
F[c･,.8 .Distribution ofnectar glands on lids collected at KP. The neetar glands of
ALepenthe svieillardii were sparsely distribuie dthroughout the entire lid ,b,
Longitudinal sections ofpitehers in KP The distributio nofthe digestive gland zone
was variable in cach pitche rT.he arro", shows the boundary between the digestive
g]and zone and the waxy zone.
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