A Monograph of Paris (Melanthiaceae): Morphology, Biology, Systematics and Taxonomy

211 Pages·2020·21.2 MB·English

by  Yunheng Ji

Yunheng Ji
A Monograph
of Paris
(Melanthiaceae)
Morphology, Biology, Systematics and Taxonomy
Paris
A Monograph of (Melanthiaceae)
Yunheng Ji
Paris
A Monograph of
(Melanthiaceae)
Morphology, Biology, Systematics
and Taxonomy
YunhengJi
KunmingInstituteofBotany
ChineseAcademyofSciences
Kunming,Yunnan,China
In association with: Zhenyan Yang, Jin Yang, Lingling Yu, Haiyang Liu, Junbo Yang, Lifang Yang,
ChangkunLiu,MinShen,LeiJinandShuyingWang.
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Dedicatedtomyprofessors,HengLi,ShengjiPei,andZhekunZhou,aswellas
to my family.
Preface
ParisLinn.isamorphologicallydistinctivegenusbelongingtothemonocotyledonousfamily
Melanthiaceae(Liliales),accordingtotheupdatedclassificationoftheAngiospermPhylogeny
Group.SpeciesofthisgenusarewidelydistributedthroughoutEurasia,enteringtheArcticat
the northernmost limit and reaching Hainan Island and Indochina at the southernmost limit.
Nearly all Paris species with a thick rhizome (medicinal Paris) are traditionally used as
medicinal herbs in China, Bhutan, India, Myanmar, Nepal, and Vietnam, owing to their
analgesic, hemostatic, anti-tumor, and anti-inflammatory activities. To date, approximately
90 commercial drugs and health products have been developed in China using the rhizomes
of medicinal Paris as raw materials. It is estimated that the income generated by the sale of
thesedrugsandproductsisapproximately10billionCNY(ca.1.6billionUSD)peryear.
DespitethegreateconomicimportanceoftheParisspecies,thetaxonomyofthegenusParis
remains controversial. First, the circumscription of this genus, long in dispute, has not been
satisfactorily resolved. Historically, Paris was recognized as either a single genus or divided
intothreenarrowlydefinedgenera(Daiswa,Kinugasa,andPariss.s.)bydifferentresearchers.
Similarly,previousphylogeneticanalysesofParisbasedondifferentDNAsequencedatasets
have yielded conflicting results. Second, the infrageneric classification of Paris is still ques-
tionable and controversial. To date, five infrageneric systems have been outlined, in which a
totaloftwosubgeneraandeightsectionshavebeenrecognized.Itremainsunclearhowmany
subgeneraandsectionsshouldberecognizedwithinthisgenus,andwhataretheevolutionary
relationships among them. Third, the taxonomy of Paris is very complicated because of the
diversity and plasticity of morphological characteristics at the species level. Approximately
95 taxa have been described to date. Although four major taxonomical revisions have been
doneonParissince1888,theyaregreatlycontradictoryindelineatingspecies.Asaresult,the
alpha taxonomy of Paris remains uncertain. To understand the true species diversity in this
genus,acomprehensivetaxonomicrevisionisurgentlyneeded.
Under the supervision of Prof. Heng Li, who has carried out the most recent and compre-
hensive taxonomic revision on this genus, my monographic study on the genus Paris was
initiated in 2002 when I was a Ph.D. student. During the past 18 years, I have conducted
extensive field observations and collections in China and neighboring countries and have
examinedthousandsofspecimens(andspecimenphotos)from88herbariaworldwide.Ihave
investigated the general morphology, pollen morphology, and genome size of Paris species.
Withtheadventofnext-generationDNAsequencingtechnologies,Igeneratedcompleteplastid
genomesandnuclearribosomalDNAsequences,toreconstructphylogeniesandtoassessthe
speciesdelineationinthisgenus.Consequently,mostofthetaxonomicissuesmentionedabove
have beensatisfactorily elucidated. This monograph outlines arevisedinfragenericsystemof
Paristhatincludesfivesections.Basedoninfragenericevolutionaryrelationshipsandmolecu-
lardating,thebiogeographicscenarioofgenusParisisproposed.
Thisbook,whichincludessevenchapters,providesessentialknowledgeonthetaxonomic
history, morphology, biology, economic importance, phytochemistry, pharmaceutical
vii
viii Preface
prospects,origin,evolution,lineagediversification,biogeography,taxonomy,andconservation
of the genus Paris, as well as the description, characteristics, illustration, and distribution of
each species. Given the great economic importance of this genus, such knowledge will be
conducivetotheexplorationandprotectionofParisspecies.
Kunming,China YunhengJi
December12,2019
Acknowledgements
This monographic study of Paris was jointly supported by the National Natural Science
Foundation of China (30670312, 31070297, 31770391, and 31872683); the NSFC-Joint
FoundationofYunnanProvince(U1802287);theMajorProgramofNationalNaturalScience
Foundation of China (31590820 and 31590823); a grant from the Large-Scale Scientific
Facilities of the Chinese Academy of Sciences (No. 2017-LSF-GBOWS-02); and the open
research project of “Cross-Cooperative Team” of the Germplasm Bank of Wild Species,
KunmingInstituteofBotany,ChineseAcademyofSciences.
The present work was accomplished with the assistance of many people. First, I should
expressmysincerethankstoChengjinYang,GuohuaZhou,RongLi,ZhilingDao,SirongYi,
QiliangGan,TingzhouZhao,RenZhao,ZhangmingWang,YongxingXiong,TaiSu,Yuling
Huang, Xiaochun Shi, Qiuping Tan, Liangxue Sun, Shaotian Chen, Wenyun Chen, Tao Su,
YongjiangHuang,YingYang,YanhongZhai,HaichengAn,YuhuaLi,YulongLi,andBenxi
Liufortheirinvaluablehelpwiththefieldobservationsandspecimencollections.Mysincere
thanks also go out to Royal Botanic Garden (Edinburg), P. Bruggeman, J. Marata,
J. McClments, and W. B. Zomlefer for providing DNA samples of Paris and related plant
taxa.DeservingspecialthanksareZhirongZhang,JingYang,andZhengshanHefortheirhelp
with the molecular experiments, Drs. Jianjun Jin, Jiahui Chen, and Hongtao Li for their
assistanceindataanalysis,ZhijiaGuforhelpingwiththemicroscopicinvestigationofpollen
morphology,andYanxiaJiaformeasuringthegenomesizesofParisspecies.Iwouldliketo
thankallstaffattheherbariaBJM,CDBI,HIB,HITBC,HWA,IBK,IGA,IMC,IMDY,KUN,
LBG, LZU, NAS, PE, SYS, SZ, WUK, and YUKU for accommodating my examination of
their specimens and to curators of the herbaria B, BN, E, GH, K, and MO for providing
specimensonloanorspecimenphotographs.Finally,duringthelast10years,mystudentshave
madegreatcontributionstothisbook,withouttheirassistanceandcomments,thisbookwould
neverhavebeenwritten.
ix
Contents
1 Introduction:TaxonomicHistory. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1 EstablishmentoftheGenus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Affinities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.3 HistoryofSpeciesDiscoveries. . . . . . . .. . . . . . . . . . . .. . . . . . . . . . . .. 3
1.4 GenericCircumscriptionandSubdivision. . . . . . . . . . . . . . . . . . . . . . . . . 8
1.5 IssuestoBeResolved. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2 Morphology.. . . .. . . . .. . . .. . . .. . . . .. . . .. . . .. . . . .. . . .. . . .. . . . .. 13
2.1 GrossMorphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
2.1.1 Rhizome. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
2.1.2 Mycorrhiza. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.1.3 Leaves. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.1.4 Flower. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.1.5 Fruit. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.1.6 Seeds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.1.7 TaxonomicSignificance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.2 PollenMorphology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
2.2.1 PollenSize,Shape,andOrnamentation. . . . . . . . . . . . . . . . . . . . . 19
2.2.2 TaxonomicImplications. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
3 Biology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3.1 DistributionandHabitat. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3.2 Phenology. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
3.3 PropagationandGrowth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
3.3.1 Pollination. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
3.3.2 SeedProduction. . . . .. . . . . . . . . . . . . . .. . . . . . . . . . . . . . .. . . 35
3.3.3 SeedDispersal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
3.3.4 LifeCycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
3.4 Chromosomes. . .. . . . .. . . . .. . . . .. . . . .. . . . . .. . . . .. . . . .. . . . .. 36
3.4.1 Karyotype. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.4.2 BChromosomes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.4.3 ChromosomeEvolution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
3.5 Genomes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
3.5.1 GenomeSize. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
3.5.2 PossibleMechanismsUnderlyingtheFormationofLarge
Genomes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
3.5.3 PlastidGenomes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
xi
xii Contents
4 EconomicImportance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
4.1 MedicinalImportance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
4.1.1 OverallInformation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
4.1.2 TraditionalUsesandEthnopharmacologicalProperties. . . . . . . . . . 55
4.2 HorticulturalPotential. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
4.2.1 P.cronquistii(Takht.)H.Li. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
4.2.2 P.delavayiFranch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
4.2.3 P.lancifoliaHayata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
4.2.4 P.luquanensisH.Li. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
4.2.5 P.marmorataStearn. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
4.2.6 P.yunnanensisFranch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
4.3 Phytochemistry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
4.3.1 SteroidalSaponins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
4.3.2 Phytoecdysones. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
4.3.3 Phytosterols. . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . 63
4.3.4 Flavonoids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
4.3.5 TriterpenoidSaponins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
4.3.6 Others. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
4.4 PharmaceuticalProspects. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
4.4.1 Anti-Tumor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
4.4.2 HemostaticActivity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
4.4.3 UterineContractileAgonisticActivity. . . . . . . . . . . . . . . . . . . . . . 65
4.4.4 Antimicrobial. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
4.4.5 OtherActivities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
5 Phylogeny,Classification,Biogeography,andEvolution. . . . . . . . . . . . . . . . 69
5.1 AncientandRecentHybridization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
5.2 GenericCircumscription. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
5.3 InfragenericClassification. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
5.4 HistoricalBiogeography. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
5.5 SpeciesDiversification. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
5.6 OriginandEvolutionofGenomicGigantism. . . . . . . . . . . . . . . . . . . . . . 77
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
6 TaxonomicRevision. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
6.1 SectionParis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
6.2 SectionKinugasa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
6.3 SectionThibeticae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
6.4 SectionAxiparis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
6.5 SectionEuthyra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
7 ConservationConsiderations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
7.1 ConservationStatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
7.1.1 SpeciesRarity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
7.1.2 ExtinctionRisk. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
7.2 PrincipalThreats. . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . 195
7.2.1 CommercialCollecting. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
7.2.2 ArbitraryIntroduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
7.2.3 HabitatDegradation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
7.3 ConservationPriorities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
7.4 ConservationStrategies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199