Saururaceae - Species Plantarum Programme

Saururaceae - Species Plantarum Programme

AUSTRALIAN BIOLOGICAL RESOURCES STUDY, CANBERRA
SPECIES PLANTARUM
FLORA OF THE WORLD
Part 11. SAURURACEAE
by Anthony R. Brach & Xia Nian-he
© Commonwealth of Australia 2005
This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be
reproduced or distributed by any process or stored in any retrieval system or data base without prior
written permission from the copyright holder. Requests and inquiries concerning reproduction and rights
should be addressed to the Director, Australian Biological Resources Study, GPO Box 787, Canberra,
ACT 2601, Australia.
EDITORS
Annette J.G. Wilson
Anthony E. Orchard
ASSISTANT EDITOR
Brigitte Kuchlmayr
This work may be cited as:
This book is available from:
Anthony R. Brach & Xia Nian-he, Saururaceae,
Species Plantarum: Flora of the World Part 11:
1–12 (2005).
Australian Biological Resources Study
GPO Box 787
Canberra ACT 2601
Australia
National Library of Australia
Cataloguing in Publication entry
Brach, Anthony R.
Saururaceae.
Includes index.
ISBN 0 642 56841 3.
1. Saururaceae. I. Xia, Nian-he. II. Australian Biological Resources Study.
III. Title. (Series : Species plantarum: flora of the world ; pt. 11).
583.25
CONTENTS
Contributors
Species Plantarum Committee
Introduction
Acknowledgments
Saururaceae
iv
v
vii
viii
1
Saururus
2
Gymnotheca
5
Anemopsis
6
Houttuynia
7
Maps
10
Index
12
iii
CONTRIBUTORS TO THIS PART
Text
Anthony R. Brach, Missouri Botanical Garden, c/o Harvard University Herbaria, 22 Divinity
Avenue, Cambridge, MA 02138-2094, United States of America
Xia, Nian-he, Herbarium IBSC, Department of Taxonomy, South China Institute of Botany,
Chinese Academy of Sciences, Wushan, Guangzhou, Guangdong 510650, China
Illustrations
Illustrations in this work have been reproduced from Z.Y.Wu, & P.H.Raven (eds.), Fl. China,
Illustr. 4: 122 (2001), with kind permission from Missouri Botanical Garden Press.
iv
Species Plantarum Steering Committee (as at May 2005)
G. Achoundong, Herbier National de Cameroon, Yaoundé, Cameroon
C.E. Anderson, University of Michigan, Ann Arbor, USA
A. Anton, Museo Botánico, Universidad Nacional de Córdoba, Argentina
W. Berendsohn, Botanischer Garten und Botanisches Museum, Berlin, Germany
D.E. Boufford, Harvard University Herbaria, Cambridge, Mass., USA
L. Boulos, Dokki, Cairo, Egypt
R.K. Brummitt, Herbarium, Royal Botanic Gardens, Kew, UK
S. Castroviejo, Real Jardín Botánico, CSIC, Madrid, Spain
A.C. Chikuni, Royal Norwegian Embassy, Lilongwe, Malawi
M.D. Correa, Universidad de Panamà, Panama
T.F. Daniel, California Academy of Sciences, San Francisco, Calif., USA
D.Darnaedi, Kebun Raya Indonesia, Bogor, Indonesia
P. Dávila, Escuela Nacional de Estudios Profesionales, UNAM, Tlalnepantla, Mexico
T.S. Filgueiras, Reserva Ecológica do IBGE, Brasilia, Brazil
E. Forero, Universidad Nacional de Colombia, Bogotá, Colombia
I. Friis, Botanical Museum and Library, Copenhagen, Denmark
V. Funk, Smithsonian Institution, Washington DC, USA
D. Geltman, Komarov Botanical Institute, St Petersberg, Russia
A.S. George, Kardinya, WA, Australia
W. Greuter, Botanischer Garten und Botanisches Museum, Berlin, Germany
C. Hilton-Taylor, I.U.C.N., Cambridge, UK
Hong D.Y., Chinese Academy of Sciences, Beijing, China
K. Iwatsuki, Yokohama, Japan
B. Jonsell, Uppsala, Sweden
B. Khayota, National Museums of Kenya, Nairobi, Kenya
J. Kirschner, Academy of Sciences, Průhonice, Czech Republic
H.P. Linder, Institut für Systematische Botanik, Zurich, Switzerland
P.-A. Loizeau, Conservatoire et Jardin Botaniques, Genève, Switzerland
J.L. Luteyn, New York Botanic Gardens, Bronx, New York, USA
K. Marhold, Slovak Academy of Sciences, Bratislava, Slovakia
M. Moraes, Universidad Mayor de San Andres, La Paz, Bolivia
P. Morat, Muséum National d'Histoire Naturelle, Paris, France
N.R. Morin, The Arboretum at Flagstaff, Arizona, USA
Nguyen T.H., Institute of Ecology & Biological Resources, Hanoi, Vietnam
H. Ohashi, Tohoku University, Sendai, Japan
R.J. Pankhurst, Royal Botanic Garden, Edinburgh, UK
M. Sanjappa, Botanical Survey of India, Kolkata, India
T. Santisuk, Royal Forest Department, Bangkok, Thailand
Sebsebe Demissew, Addis Ababa University, Addis Ababa, Ethiopia
J.H. Seyani, National Herbarium and Botanic Gardens, Zomba, Malawi
G.F. Smith, National Botanical Institute, Pretoria, South Africa
W.D. Stevens, Missouri Botanical Garden, Saint Louis, Missouri, USA
v
W.L. Wagner, Smithsonian Institution, Washington DC, USA
A.J.G. Wilson, Australian Biological Resources Study, Canberra, ACT, Australia
K. Wilson, Royal Botanic Gardens, Sydney, NSW, Australia
vi
INTRODUCTION
Species Plantarum aims to provide in concise format, and with standardised data fields, basic
taxonomic information on the vascular plants of the world, including accepted names and
synonyms with bibliographic data, types of names, keys and descriptions from family to
varietal levels, geographical distributions, ecological information and other related matters,
and to publish it in both hard copy and electronic form.
The format of the Species Plantarum is based on that of Flora of Australia, with some
departures made necessary by the different scale of the project. Initially at least, the series is
being edited and published for the Species Plantarum Project and IOPI by the Australian
Biological Resources Study (ABRS), producers of Flora of Australia.
Treatments are contributed on a voluntary basis. Each part of Species Plantarum is intended
to provide a complete account of a family, subfamily, large genus or other related taxonomic
group. While treatments of small families may be shorter, it is intended that contributions
will, in general, cover at least 50 to 100 species. The taxonomy adopted is that of the author,
although the family delimitations recommended are initially those of R.K.Brummitt,
Vascular Plant Families and Genera (1992). The order of taxa within families, genera and
species in the Species Plantarum is intended to reflect natural relationships, so far as this is
possible in a linear sequence.
Maps are provided for each species, or in those cases where infraspecific taxa are recognised,
for each of the terminal taxa. Distribution maps are based on those in R.K.Brummitt, World
Geographical Scheme for Recording Plant Distributions 2nd edn (2001), and the ‘countries’
adopted are those of Level 3 and 4 of that work. Description of distribution follows the same
work, with a two-digit code for regions and a three-letter code for the 'country'. Upper case
letters for the 'country' indicate native distribution; lower case letters indicate that the taxon
is only present in that 'country' as an introduced and naturalised plant. If a taxon is extinct in
a 'country', this is indicated by a dagger. Distribution of species as cultivated plants is not
included.
Misapplied and invalid names are, in general, omitted. Journal titles are abbreviated
according to G.D.R.Bridson & E.R.Smith, Botanico-Periodicum-Huntianum / Supplementum
(1991). Book titles are abbreviated according to F.A.Stafleu & R.S.Cowan, Taxonomic
Literature (2nd edn) Vols 1–7, and Supplements (1976-), except that upper case initial letters
are used for proper names and significant words. Authors of plant names are abbreviated
according to R.K.Brummitt & C.E.Powell, Authors of Plant Names (1992).
A separate part, Introduction to the Series, provides a history of the project, a glossary,
guide for contributors and key to the conventions used in describing distribution. These
resources will also be available on the World Wide Web, initially through the ABRS site
(currently http://www.deh.gov.au/biodiversity/abrs) with links from the IOPI site (currently
http://plantnet.rbgsyd.gov.au/iopi/iopihome.htm) and others.
A.J.G.Wilson
Canberra
May 2005
vii
ACKNOWLEDGMENTS
The authors wish to thank Shirley C. Tucker for helpful review comments, and
Kanchi N. Gandhi and Jan Kirschner for nomenclatural advice.
viii
SAURURACEAE
Anthony R. Brach & Xia Nian-he
Saururaceae Rich. ex E.Mey., Houttuynia 20 (1827), nom. cons.
Type: Saururus L.
Herbs perennial, aromatic. Stems erect, ascending or prostrate, obviously jointed. Leaves
alternate, simple; stipules interpetiolately connate or adnate to petiole, forming a sheath.
Inflorescence a dense spike or raceme, sometimes resembling a single flower when large
involucral petaloid bracts are present at base. Flowers bisexual; perianth absent. Stamens
usually 3, 6, or 8, free or adnate to ovary; anthers 2-locular, longitudinally dehiscent. Pistils
(2 or) 3 or 4-carpellate, rarely to 7-carpellate; carpels distinct or connate, if distinct then
each carpel with 2–4 ovules, if connate then ovary unilocular, with placentation parietal and
each placenta with 6–13 ovules; styles free. Fruit a schizocarp or apically dehiscent capsule.
Seed(s) 1 or many; endosperm scanty; perisperm abundant; embryo minute.
Four genera, six species, in temperate to tropical regions of S & E Asia and N America; three
genera and four species in Asia; two genera each having one species in N America.
The species of this family have been extensively compared in studies of floral vasculature
(H.X.Liang & S.C.Tucker, Amer. J. Bot. 77: 607–623, 1990), organogenesis (H.X.Liang,
Acta Phytotax. Sin. 32: 425–432, 1994), pollen morphology (H.X.Liang, Acta Bot. Yunnan.
14: 401–404, 1992), and stem anatomy (S.Carlquist et al., Internatl Assoc. Wood Anat. J.
16(2): 133–150, 1995). Oil cells are present throughout Saururaceae, and are the basis for the
"aromatic" nature of both vegetative and reproductive organs in the family, and have been
studied developmentally (S.C.Tucker, Bot. Gaz. 137: 341–347, 1976). Phylogeny within the
family (S.W.Meng et al., J. Pl. Res. 115: 71–76, 2002), with its sister clade Piperaceae
(S.C.Tucker et al., Syst. Bot. 18: 614–641, 1993), and another genus of early angiosperms —
Nymphaea (J.-F.Leroy, Evolution 319: 543–547, 1996) has been discussed. Initially included
in the Saururaceae, the genus Circaeocarpus C.Y.Wu (Acta Phytotax. Sin. 6(2): 222, 1957) is
treated as a member of the Piperaceae (= Zippelia Blume, see Y.Q.Cheng et al., Fl. China
4: 110, 1999).
G.F.Buddell & J.W.Thieret, Saururaceae, Fl. N. America N. Mexico 3, Magnoliophyta:
Magnoliidae and Hamamelidae, 37–38 (1997); Y.C.Tseng, Saururaceae, in Y.C.Tseng (ed.),
Fl. Reipubl. Popularis Sin. 20(1): 4–11 (1982); S.C.Tucker et al., Utility of onto-genetic and
conventional characters in determining phylogenetic relationships of Saururaceae and
Piperaceae (Piperales), Syst. Bot. 18: 614–641 (1993); C.E.Wood, Jr., The Saururaceae in the
southeastern United States, J. Arnold Arbor. 52: 479–485 (1971); N.H.Xia & A.R.Brach,
Saururaceae, in Z.Y.Wu & P.H.Raven (eds.), Fl. China 4: 108–109 (1999).
KEY TO GENERA
1
Herbs prostrate, ±stoloniferous; stamens shorter than styles
1:
2. GYMNOTHECA
Herbs erect or ascending, rhizomatous; stamens longer than styles
2
Inflorescence a raceme, without involucral petal-like bracts; flowers
with elongate flower-bract stalk (“pedicel”); fruit a schizocarp
2:
3
3:
1. SAURURUS
Inflorescence a dense spike, with 4 or more involucral petal-like bracts
at base; flowers sessile; fruit a capsule
Basal leaves present; basal lamina elliptic-oblong; cauline leaves
dimorphic; stamens 6 (–8); inflorescence with 4–9 involucral bracts at
base (North America)
Basal leaves absent; cauline lamina broadly ovate or ovate-cordate;
stamens 3, rarely 4; inflorescence with 4 (rarely 6 or 8) involucral
bracts at base (Asia)
3. ANEMOPSIS
4. HOUTTUYNIA
1
SPECIES PLANTARUM —FLORA OF THE WORLD
2005
1. SAURURUS
Saururus L., Sp. Pl. 1: 341 (1753)
Type: Saururus cernuus L.
Spathium Lour., Fl. Cochinch. 217 (1790).
T: Spathium chinense Lour.
Mattuschkia J.F.Gmel., Syst. Nat., 13th edn [bis] 2: 589 (1791), non Mattuschkaea Schreb. (1791).
T: Mattuschkia aquatica (Walter) J.F.Gmel.
Saururopsis Turcz., Bull. Soc. Imp. Naturalistes Moscou 21(1): 589 (1848).
(Lour.) Turcz.
T: Saururopsis chinensis
Herbs, erect or ascending, rhizomatous. Stems longitudinally ridged and sulcate. Leaves:
stipules membranous; petiole shorter than lamina; lamina entire, with 5–7 basal nerves.
Inflorescence a raceme, leaf-opposed and/or terminal, without involucral bracts; rachis
pubescent. Flowers 100–350, small, white when mature, with elongate flower-bract stalk
(“pedicel” see Liang & Tucker, 1990). Stamens usually 6, sometimes 8, rarely 3, free, longer
than styles; filaments c. as long as or slightly longer than anthers; anthers oblong. Pistils 4 or
1, 3 or 4 (–7)-carpellate, with carpels distinct or connate at base; ovary superior; each carpel
with 2–4 ovules; styles 4 or 1, recurved. Fruit a schizocarp; mericarps 3 or 4, indehiscent.
Seed 1 per mericarp.
Two species, distributed disjunctly in E North America and E Asia.
T.Holm, Saururus cernuus L. A morphological study, Amer. J. Sci., ser. 5, 12: 162–168
(1926); S.C.Tucker, Floral development in Saururus cernuus (Saururaceae). 1. Floral
initiation and stamen development, Amer. J. Bot. 62: 993–1007 (1975); S.C.Tucker, Floral
development in Saururus cernuus (Saururaceae). 2. Carpel initiation and floral vasculature,
Amer. J. Bot. 63: 289–301 (1976); S.C.Tucker, Ontogeny of the inflorescence of Saururus
cernuus (Saururaceae), Amer. J. Bot. 66: 227–236 (1979).
Racemes nodding to erect; floral bracts green; leaf petioles to 10 cm long;
stamens hypogynous, filaments 3–4 mm long (North America)
Racemes not nodding, floral bracts white; leaf petioles to 3 cm long; stamens
epigynous, adnate to lower part of ovaries, filaments c. 1 mm long (Asia)
1. S. cernuus
2. S. chinensis
1. Saururus cernuus L., Sp. Pl. 1: 341 (1753)
T: [U.S.A., Virginia], "Habitat in Marilandia, Virginia"; lecto: Herb. Linn. 478.1, LINN, fide J.L.Reveal et al.,
Huntia 7: 234 (1987).
Mattuschkia aquatica (Walter) J.F.Gmel., Syst. Nat., 13th edn [bis] 589 (1791); Saururus aquaticus Walter,
Fl. Carol. 127 (1788). T: not designated.
Saururus lucidus Donn, Hortus Cantabrig., 3rd edn 66 (1804).
T: [U.S.A.], Sep. 1791, s.coll.; holo: n.v.
Saururus cernuus L. f. submersus Glück, Biol. Morphol. Untersuch. Wasser-Sumpfgewächse 3: 69, t. 3 (1911).
T: not designated, cultivated.
Illustrations: H.X.Liang & S.C.Tucker, Amer. J. Bot. 77: 609, 612 (1990); L.Watson & M.J.Dallwitz, Fam. Fl.
Pl., http://delta-intkey.com (1992–).
Map: G.F.Buddell & J.W.Thieret, Saururaceae, in Fl. N. America N. Mexico 3, Magnoliophyta: Magnoliidae
and Hamamelidae 38 (1997).
Herbs 15–120 cm high, mostly pubescent when young, often glabrate; rhizome extensive,
stout, whitish, often with adventitious roots. Leaves: stipule sheath 2–10 mm long, slightly
clasping; petiole 1–10 cm long; lamina ovate or cordate, 2–18 × 1–10 (–15) cm, cordate
at base, acuminate, papery, with 5–7 nerves. Racemes terminal and/or leaf-opposed, nodding
to erect, narrow, (4–) 5–35 cm long; peduncle 2–15 mm long. Bract beneath each flower
adnate to pedicel, navicular, 1.5–3 mm long (distinct portion), green. Stamens hypogynous,
3–4 mm long. Fruit 1.5–3 mm long, rugose. 2n = 22, fide H.Okada, Bot. Mag. (Tokyo) 99:
289–299 (1986).
Eastern Canada to C and E USA; introduced in Germany and Italy. 11: GER. 13: ITA.
72: ONT, QUE. 74: ILL, KAN, MSO, OKL. 75: CNT, INI, MIC, NWJ, NWY, OHI, PEN,
2
SAURURACEAE (Saururus)
Figure 1. Saururus chinensis (Lour.) Baill. A, lower part of stem; B, flowering branchlet;
C, flower; D, schizocarp; E, mericarp. Scale bars: A, B = 2 cm; C–E = 2 mm. Drawn by
Deng Yingfeng & Yu Hanping. Reproduced from Z.Y.Wu & P.H.Raven (eds.), Fl. China,
Illustr. 4: 121 (2001), with permission from Missouri Botanical Garden Press.
3
SPECIES PLANTARUM —FLORA OF THE WORLD
2005
RHO, WVA. 77: TEX. 78: ALA, ARK, DEL, FLA, GEO, KTY, LOU, MRY, MSI, NCA,
SCA, TEN, VRG. Swamps, marshes, ponds, slow-moving streams, river banks, wet soil,
fresh or slightly brackish water to 50 cm depth. Flowering (Feb.–) Mar.–Aug. (–Sep.);
fruiting Jun.–Oct. (–Nov.). Map 1.
72. ONTARIO: edge of Grand R., F.H.Montgomery 557 (GH). QUEBEC: Sainte-Rose, F.Marie-Victorin &
F.Rolland-Germain 55107 (MO).
74. ILLINOIS: St. Claire Co., J.O.Neill 15954 (MO). MISSOURI: E of
Eudora, Polk Co., J.A.Steyermark 24058 (MO).
75. PENNSYLVANIA: Dauphin Co., Jun Wen 1100 (GH).
78. ARKANSAS: Lee Co., D.Demarée 12979 (MO, GH). FLORIDA: Nassau Co., R.K.Godfrey & W.Lindsey
56849 (GH). NORTH CAROLINA: Carteret, J.Stone & D.Holland 1515 (MO). SOUTH CAROLINA:
Charleston Co., S.R.Hill 23825 (MO, GH). VIRGINIA: Miners Pond, Prince George Co., M.L.Fernald &
B.Long 8227 (GH).
Some Native Americans used this species medicinally (D.E.Moerman, Med. Pl. Nat.
America, 1986; D.E.Moerman, Nat. Amer. Ethnobot., 1998).
2. Saururus chinensis (Lour.) Baill., Adansonia 10(2): 71 (1871)
Spathium chinense Lour., Fl. Cochinch. 217 (1790); Saururus loureiri Decne., Ann. Sci. Nat., Bot., sér. 3,
3: 102 (1845), nom. illeg.; Saururopsis chinensis (Lour.) Turcz., Bull. Soc. Imp. Naturalistes Moscou
21: 590 (1848). T: [China] "Habitat in locis paludosis, prope Cantonem Sinarum", J. de Loureiro 2171; holo: P.
Saururopsis cumingii C.DC., in A. de Candolle, Prodr. 16: 239 (1869).
H.Cuming 1259; holo: BM.
T: [Philippines] In ins. Philippinis,
Saururopsis cumingii var. japonica C.DC., in A. de Candolle, Prodr. 16: 240 (1869). T: [Japan] In Japonia ad
Kanagavia, R.Fortune; syn: BM; [Japan] In Japonia ad Kanagavia, Nagasahi, R.Oldham 668; syn: G-DC, n.v.
[Saururus cernuus auct. non L.: C.P.Thunberg, Fl. Jap. 154 (1784)]
Saururus sinensis Teijsm. & Binn., Cat. Hort. Bot. Bogor. 77 (1866), nom. inval., probably an orthographic
variant of the name S. chinensis.
Illustrations: T.S.Liu & H.C.Wang (eds.), Fl. Taiwan 2: 623 (1996); K.M.Liu, Fl. Hunan. 2: 787 (2000);
Z.Y.Wu & P.H.Raven (eds.), Fl. China, Illustr. 4: 121 (2001).
Maps: H.L.Li, Trans. Amer. Philos. Soc. n.s. 42(2): 411 (1952); L.K.Fu & T.Hong (eds.), Higher Pl. China 3:
316 (1999).
Herbs to more than 1 m high; rhizomes stout, creeping, white. Leaves: stipular sheath 2–10
mm long, slightly clasping; petiole 1–3 cm long, glabrous; lamina ovate to ovate-lanceolate,
(4–) 10–20 × (2–) 5–10 cm, cordate or oblique-cordate at base, acute or acuminate at apex,
papery, densely glandular, glabrous; nerves 5–7, basal, if 7-nerved, then outermost pair
slender, ascending and then arched and combined. Apical leaves smaller, 2 or 3 at stem apex,
usually petal-like, white at anthesis. Inflorescence an elongated leaf-opposed and/or terminal
raceme, (3–) 12–20 (–22) cm long; peduncle 0.5–4.5 cm long, glabrous. Bract beneath each
flower spathulate, 1–1.5 mm long (distinct portion), white; basal part linear and pilose;
apical part small, orbicular, rounded to acute at apex, glabrous or sparsely ciliate. Stamens
epigynous, adnate to lower part of ovaries; filaments slightly longer than anthers, c. 1 mm
long. Fruit c. 3 mm diam., tuberculate. 2n = 22, fide C.J.Ge et al., in D.Hong (ed.),
Pl. Chromosome Res. 1987: 267–272 (1989). Fig. 1.
China, Eastern Asia, Indo-China, and Malesia. 36: CHC, CHH, CHS. 38: JAP, KOR, TAI.
40: IND. 41: VIE. 42: PHI. Wet places, ditches, meadows, riverbanks, streamsides, forests,
thickets, roadsides, field margins. Flowering Apr.–Jun., fruiting Jun.–Jul. Map 2.
36. CHINA SOUTHEAST: Iu Village, Lingnan, Guangdong, To & Ts'ang 12113 (MO); Longzhou, Guangxi,
S.P.Ko 55156 (IBSC); Liping, Guizhou, Y.K.Li 9101 (IBSC); Yang Shan, Changning Xian, Hunan, C.S.Fan &
Y.Y.Li 297 (BM, A); Anyuan, Jiangxi, C.M.Hu 2566 (IBSC); Hong Kong, S.Y.Hu & P.Y.But 20777 (A).
CHINA SOUTH-CENTRAL: Xingshan, Hubei, G.X.Fu & Z.S.Zhang 213 (IBSC); Emei Shan, Sichuan,
W.P.Fang 18901 (A); W Tianmushan, M.B.Deng 4321 (IBSC).
38. TAIWAN: Taipei, T.I.Chuang &
M.T.Kao 3378 (A).
In China, the rhizomes and flowers are used medicinally. In India, the rhizomes are boiled
and eaten.
4
SAURURACEAE (Saururus)
Excluded name
Saururus natans L., Mant. Pl. 2: 227 (1767–1771)
This is Aponogeton natans (L.) Engl. & K.Krause, in H.G.A.Engler, Pflanzenr., Aponogetonac.
11 (1906).
2. GYMNOTHECA
Gymnotheca Decne., Ann. Sci. Nat., Bot., sér. 3, 3: 100 (1845)
Type: Gymnotheca chinensis Decne.
Herbs, prostrate, ±stoloniferous, glabrous. Stems longitudinally ridged and sulcate, 30–70 cm
long, rooted at nodes. Leaves: stipules membranous, enlarged and clasping at base, slender;
petiole nearly as long as or longer than lamina; lamina papery, subglabrous, ±glandular,
entire or inconspicuously crenulate; nerves 5–7, basal, with outermost pair of nerves sometimes slender or inconspicuous. Inflorescence a leaf-opposed and/or terminal spike, with large
involucral white leafy bracts at base or not; rachis compressed, with both sides broadly
ridged or nearly winged. Flowers 10–20, small, white, with short flower-bract stalk (“pedicel”).
Stamens (5) 6 (7), adnate to and nearly on apex of ovary, shorter than styles; filaments c. as
long as to slightly longer than anthers; anthers oblong. Pistils 4, (2 or 3) 4-carpellate; carpels
connate; ovary semi-inferior, 1-locular; placentae 4, each with 9–13 ovules; styles 4,
recurved. Fruit a fusiform capsule, dehiscent at apex. Seeds 8–40.
Two species in C and SW China, and N Vietnam.
T.S.Ying et al., End. Gen. Seed Pl. China, Saururaceae, 647–649 (1993).
Inflorescences without involucral leafy bracts at base
Inflorescences with 3 or 4 large involucral white leafy bracts at base
1. G. chinensis
2. G. involucrata
1. Gymnotheca chinensis Decne., Ann. Sci. Nat., Bot., sér. 3, 3: 100 (1845)
T: [China], s. coll.; holo: P, n.v.
Saururus cavaleriei H.Lév., Repert. Spec. Nov. Regni Veg. 10: 149 (1911).
H.Léveillé; holo: P, n.v.
T: China, Guizhou (Kweichau),
Gymnotheca weinmannifolia hort. ex L.Gentil, Liste Pl. Cult. Serres Jard. Bot. Bruxelles 95 (1907), nom. nud.,
inval. T: locality not given, cultivated material.
Illustrations: J.Decaisne, Ann. Sci. Nat., Bot., sér. 3, 3: 100, fig. 4. (1845); T.S.Ying et al., End. Gen. Seed Pl.
China, Saururaceae, 648 (1993); Z.Y.Wu & P.H.Raven (eds), Fl. China, Illustr. 4: 122 (2001).
Maps: T.S.Ying et al., End. Gen. Seed Pl. China, Saururaceae, 649 (1993); L.K.Fu & T.Hong (eds.), Higher
Pl. China 3: 318 (1999).
Stems usually 30–65 cm long. Leaves: stipular sheath 1.5–2 cm long, c. 1/3 as long as petiole;
petiole 1.5–7 cm long; lamina reniform-cordate, (1.5–) 3–6.5 × (3–) 4–7.5 cm, auriculate at
base, broadly acute or rounded at apex. Inflorescence 2–7.5 cm long; peduncle c. as long as
or slightly shorter than inflorescence, 3–6 cm long; involucral bracts absent. Bract beneath
each flower oblanceolate, c. 3 mm long, acute, sometimes lowermost bracts slightly larger and
nearly ligulate. Filaments nearly as long as or slightly longer than anthers. Ovary narrowly
obovoid. 2n = 18, fide H.X.Liang, Acta Bot. Yunnan. 13(3): 303–307 (1991). Fig. 2D–F.
S China to Indo-China. 36: CHC, CHS. 41: VIE. Streamsides, valleys. Flowering Apr.–Nov.
Map 3.
36. CHINA SOUTHEAST: Ruyuan, Guangdong, S.P.Ko 52965 (IBSC); Lingle Xian, Guangxi, C.Wang 43107
(MO, IBSC). CHINA SOUTH-CENTRAL: Hubei, A.Henry 4180 (BM); Guizhou, H.R.E.Handel-Mazzetti 2094
(E); Guiding, Guizhou, S.W.Teng 1643 (IBSC); Guizhou, S.H.Chun 2221 (PE); Sichou Xian (Si-chour-hsien),
Fadou (Faa-doou), Yunnan, K.M.Feng 12140 (A).
41. INDO-CHINA: Tonkin, P.A.Pételot 6298 (A);
Kompong Leeong (Leach), Purset, Cambodge, M.A.Martin 631 (P).
Used medicinally.
5
SPECIES PLANTARUM —FLORA OF THE WORLD
2005
2. Gymnotheca involucrata C.P'ei, Contr. Biol. Lab. Chin. Assoc. Advancem. Sci., Sect.
Bot. 9: 111 (1934)
T: China, Sichuan, Mapien-hsien [Mabian Xian], Ta-chu-pao, 19 May 1930, W.P.Fang 248; holo: SZ.
Illustrations: C.P'ei, Contr. Biol. Lab. Chin. Assoc. Advancem. Sci., Sect. Bot. 9: 112, fig. 11 (1934);
H.X.Liang & S.C.Tucker, Amer. J. Bot. 77: 609, 614 (1990); T.S.Ying et al., End. Gen. Seed Pl. China,
Saururaceae, 648 (1993).
Maps: T.S.Ying et al., End. Gen. Seed Pl. China, Saururaceae, 649 (1993); L.K.Fu & T.Hong (eds.), Higher
Pl. China 3: 318 (1999).
Stems usually 30–70 cm long. Leaves: stipular sheath 1.5–2 cm long, 1/4–1/3 as long as petiole;
petiole 1.5–8.5 cm long; lamina cordate or reniform-cordate, 4–18 × 6–10 cm, deeply
auriculate at base, broadly acute at apex. Inflorescence c. 5 cm long; peduncles 4–7 cm long;
involucral bracts 3 or 4, leafy, obovate-oblong, 1.2–1.8 × 0.8–1.2 cm, white. Bract beneath
each flower obovate-oblong or oblanceolate, c. 3 mm long, acute. Filaments slightly longer
than anthers. Ovary obconical. 2n = 18, fide H.X.Liang, Acta Bot. Yunnan. 13(3): 303–307
(1991).
China (S Sichuan). 36: CHC. Roadsides, wet places in forests. Flowering Feb.–Jun.
Map 4.
36. CHINA SOUTH-CENTRAL: Sichuan, E.Faber 897 (MO); Mabian, Sichuan, F.T.Wang 22225 (IBSC); Xia
Jiang, Sichuan, H.X.Liang s.n. (KUN); Sichuan, N.L.Chen 450 (KUN); Kwanhsien, Sichuan, T.P.Wang 10139
(PE); Sichuan, C.P'ei 6004 (PE); Sichuan, W.P.Fang 30859 (PE).
3. ANEMOPSIS
Anemopsis Hook. & Arn., Bot. Beechey Voy. 390 (1840)
Anemia Nutt., in J.E.Taylor, Ann. Nat. Hist. 1: 136 (1838), nom. illeg., non Anemia Sw. (1806), nom. cons.
Type: Anemopsis californica (Nutt.) Hook. & Arn.
Herbs, erect or ascending. Stems not branched; nodes 1 or 2 (–7). Leaves mostly basal; basal
petiole c. twice as long as lamina, pubescent along margin; cauline leaves sessile or
petiolate; lamina leathery, glabrous. Inflorescence a terminal spike, compact, conical,
subtended by 4–9 petaloid bracts; rachis pubescent. Flowers 75–150, small, coalescent, sessile,
epigynous. Stamens 6 (–8), adnate to ovary, longer than styles; filament c. as long as anther;
anther oblong. Pistil 1, syncarpous, 1 or 3 (or 4)-carpellate; placentae 3 (or 4); ovules 4–10
per placenta; styles and stigmas 3 (or 4), distinct. Fruit a capsule, dehiscent. Seeds 18–40.
One species, W North America.
T.Holm, Anemiopsis [sic] californica (Nutt.) H. and A. An anatomical study, Amer. J. Sci.,
ser. 4, 19: 76–82 (1905); L.Kelso, A note on Anemopsis californica, Amer. Midl. Naturalist
13: 110–113 (1932); C.H.Quibell, Floral anatomy and morphology of Anemopsis californica,
Bot. Gaz. 102: 749–758 (1941); S.C.Tucker, Initiation and development of inflorescence and
flower in Anemopsis californica (Saururaceae), Amer. J. Bot. 72: 20–31 (1985).
Anemopsis californica (Nutt.) Hook. & Arn., Bot. Beechey Voy. 390 (1840)
Anemia californica Nutt., in J.E.Taylor, Ann. Nat. Hist. 1: 136, t. 92 (1838); Houttuynia californica (Nutt.)
Benth. & Hook.f., Bot. California 2: 483 (1880).
T: [USA], California, Santa Barbara and San Diego,
s.coll.; holo: ?PH, ?BM, n.v.
Anemopsis bolanderi C.DC., Linnaea 37: 333 (1871–1873); Houttuynia bolanderi (C.DC.) Benth. & Hook.f.,
Gen. Pl. 3: 128 (1880). T: USA, California, H.N.Bolander; holo: G-DC, n.v.
Anemopsis ludovici-salvatoris Willk., in E.A. von Regel, Gartenflora 26: 228 (1877).
cultivated, s.coll.; holo: n.v.
Anemia intermedia Copel. ex M.E.Jones, Contr. W. Bot. No. 15, 123 (1929).
'El Tigre Mine', Mar. 1927, M.E.Jones 23472; holo: RSA, n.v.
T: [USA, California],
T: Mexico, Nyarit, Acaponeta,
Anemopsis californica var. subglabra Kelso, Amer. Midl. Naturalist 13(3): 112–113 (1932).
Colorado, Aurora, 'Bluff Lake' 2 mi north, 30 Jul. 1930, L.Kelso 1164; holo: US, n.v.
6
T: USA,
SAURURACEAE (Anemopsis)
Anemopsis californica var. typica Kelso, Amer. Midl. Naturalist 13(3): 111 (1932), nom. inval.
Illustrations: W.Curtis, Bot. Mag. tab. 5292 (1862); H.X.Liang & S.C.Tucker, Amer. J. Bot. 77: 609, 616 (1990).
Map: G.F.Buddell & J.W.Thieret, Saururaceae, Fl. N. America N. Mexico 3, Magnoliophyta: Magnoliidae and
Hamamelidae, 38 (1997).
Herbs, 8–80 cm long, densely pubescent to nearly glabrous, stoloniferous. Basal leaves to 14
or more; petiole 2–40 cm long; lamina elliptic-oblong, 1–25 × 1–12 cm, cordate to obtuse at
base, rounded at apex. Cauline leaves dimorphic. Primary leaf 1 (or 2) per axil, usually with
secondary leaves in axil, sessile, broadly to narrowly ovate, 1–9 × 1–4 cm, clasping at base,
rounded to acute at apex. Secondary leaves 1–4; petiole 1–12 cm long; lamina ellipticoblong, 2–10 × 1–5 cm, cordate to rounded at base, rounded to acute at apex. Spikes erect,
1–4 cm long; bracts petaloid, 5–35 × 5–15 mm, white to reddish. Floral bracts ±orbicular,
3.5–6 mm long (distinct portion), clawed, white, each adnate to an ovary. Capsules
coalescent but easily separable, 5–7 mm long, brown. 2n = 44, fide H.Okada, Bot. Mag.
(Tokyo) 99: 289–299 (1986).
Northwestern, Southcentral, and Southwestern USA to North and Central Mexico. 73: COL,
ORE, WAS. 76: ARI, CAL, NEV, UTA. 77: NWM, TEX. 79: MXN, MXE, MXS. Wet,
alkaline, saline, and coastal marsh areas. Flowering Mar.–Sep.; fruiting Apr.–Nov. Map 5.
73. COLORADO: N of Aurora, Adams Co., J.Ewan 14421, [topotype of var. subglabra] (MO, GH).
76. CALIFORNIA: San Diego, H. van der Werff 13997 (MO); near Box S Springs, Mojave Desert, San
Bernardino Co., M. & R.P.Ownbey 1682 (MO, GH). UTAH: Washington, T.G.Yuncker & W.H.Welch 4468
(MO, GH).
77. NEW MEXICO: Cuevas Canyon, San Miguel Co., S.R.Hill & P.A.Levandoski 12279 (MO, GH).
79. MEXICO NORTHWEST: Norte de Baja California, J.S.Miller, M.Merello & A.Pool 7400 (MO); Comondu,
Baja California, H.S.Gentry 4241 (MO, GH). MEXICO NORTHEAST: Durango, E.Palmer 118 (MO, GH).
Some Native Americans used the roots of this species medicinally (T.Timbrook, Econ. Bot.
44(2): 236–253, 1990).
4. HOUTTUYNIA
Houttuynia Thunb., Kongl. Vetensk. Acad. Nya Handl. 4: 149, 151 (1783), nom. cons.
T: Houttuynia cordata Thunb.
Polypara Lour., Fl. Cochinch. 34, 61 (1790).
T: Polypara cochinchinensis Lour.
Herbs, erect or ascending, rhizomatous. Stems longitudinally ridged and sulcate; basal part
of stems creeping, rooted in whorls at nodes; apical part erect, glabrous or pubescent on
nodes, sometimes purplish red. Leaves: stipules membranous; petiole shorter than lamina;
lamina entire; nerves 5–7, with basal or innermost pair arising c. 5 mm above base, if 7nerved then outermost pair very slender and inconspicuous. Inflorescence a terminal and/or
leaf-opposed densely-flowered spike, with 4 (rarely 6 or 8) involucral white petal-like bracts
at base. Flowers c. 100, small, sessile, white when mature. Stamens 3, rarely 4, longer than
styles; filaments c. 3 times as long as anthers, with basal part connate to ovary; anthers
oblong. Pistils 3, 3-carpellate; carpels partly connate; ovary semi-inferior (flowers
perigynous), unilocular; placentae 3, each with 6–9 ovules; styles 3, recurved. Fruit a
subglobose capsule, dehiscent at apex. Seeds 9–18.
One species, centred in E and S Asia, from Japan and China to the Indian Subcontinent, and
to the Malesian Region in the south; also in Madagascar.
Houttuynia cordata Thunb., Kongl. Vetensk. Acad. Nya Handl. 4: 149, 151 (1783)
Polypara cordata (Thunb.) Kuntze, Revis. Gen. Pl. 565 (1891).
C.P.Thunberg; holo: UPS, n.v.
T: [Japan] Doku Dami vulgo, it. Sjunjak,
Polypara cochinchinensis Lour., Fl. Cochinch. 34, 61 (1790).
T:
Cochinchinenses, inter olera in acetariis edulis, J. de Loureiro; holo: P, n.v.
Houttuynia foetida hort. ex Loudon, Hort. Brit. 37 (1830).
from Japan in 1800.
[Vietnam], Habitat
hortos
T: not designated, cultivated material introduced
7
SPECIES PLANTARUM —FLORA OF THE WORLD
2005
Figure 2. A–C, Houttuynia cordata Thunb. A, plant; B, inflorescence; C, flower. D–F,
Gymnotheca chinensis Decne. D, plant; E, inflorescence portion; F, longitudinal section of
flower showing ovules. Scale bars: A, B, D = 2 cm; C, E, F = 2 mm. Drawn by Huang
Shaorong. Reproduced from Z.Y.Wu, & P.H.Raven (eds.), Fl. China, Illustr. 4: 122 (2001),
with permission from Missouri Botanical Garden Press.
8
SAURURACEAE (Houttuynia)
Houttuynia cordata f. viridis J.Ohara, J. Phytogeogr. Taxon. 33(2): 72 (1985).
cho, Okazaki-shi, prov. Mikawa, 18 Jun. 1983, Sumio Honda; holo: AICH, n.v.
Houttuynia cordata f. polypetaloidea T.Yamaz., J. Jap. Bot. 61(1): 310 (1986).
Gard. cult., 30 Jun. 1986, T.Yamazaki; holo: TI, n.v.
Polypara cochinchinensis Lour., Fl. Cochinch. 34, 61 (1790).
T: [Japan] Hondo: OwhataT: [Japan] Koishikawa, Bot.
T: [Vietnam], J. de Loureiro; holo: P, n.v.
Illustrations: T.Yamazaki, Bot. Mag. (Tokyo) 91: 69–82 (1978); K.M.Liu & H.C.Wang, Fl. Taiwan 2: 621
(1996); Z.Y.Wu & P.H.Raven (eds.), Fl. China, Illustr. 4: 122 (2001).
Maps: C.Y.Wu, Acta Phytotax. Sin. 6: 189, fig. 9 (1957); L.K.Fu & T.Hong (eds.), Higher Pl. China 3: 317
(1999).
Herbs, (5–) 30–60 cm high; rhizomes creeping, thin. Leaves: stipular sheath (0.5–) 1–2.5 cm
long, 1/4–1/2 as long as petiole, usually ciliate, enlarged and slightly clasping at base; petiole
(0.7–) 1–3.5 (–4) cm long, glabrous; lamina broadly ovate or ovate-cordate, (1.5–) 4–10 ×
(1.8–) 2.5–6 cm, cordate at base, short acuminate at apex, thinly papery, densely glandular,
usually glabrous, sometimes pubescent at nerve axils, usually purplish abaxially.
Inflorescences (0.4–) 1.5–2.5 (–2.7) × (2–) 5–6 mm; peduncles 1.5–3 cm long, subglabrous;
involucral bracts oblong or obovate, (5–) 10–15 × (3–) 5–7 mm, rounded at apex. Bract
beneath each flower linear, terete, inconspicuous. Stamens longer than ovary. Capsule
2–3 mm long, with persistent styles. 2n = 96, fide H.Okada, Bot. Mag. (Tokyo) 99: 289–299
(1986). Fig. 2A–C.
West Indian Ocean, Madagascar, Indian Subcontinent, China, Eastern Asia, Indo-China,
Malesia. 29: MDG. 36: CHC, CHT. 38: JAP, KOR, NNS, TAI. 40: ASS, BAN, BHU, IND,
NEP. 41: BMA, CBD, THA. 42. JAW, SUM. Ravines, streamsides, forests, wet meadows,
slopes, thicket and field margins, trailsides, roadsides, ditch banks. Flowering Apr.–Sep.,
fruiting Jun.–Oct. Map 6.
29. MADAGASCAR: Madagascar, L.J.Dorr 2816 (MO). 36. CHINA NORTH-CENTRAL: Wenxian, Gansu,
J.X.Yang 3704 (IBSC). CHINA SOUTHEAST: Hua'an, Fujian, K.M.Wu 60118 (IBSC); Wahshan Kuek, N.T.,
Hong Kong, S.Y.Hu 10868 (A); Iu Village, Guangdong, To & Ts'ang 12077 (MO); Shap Man, Taai Shan,
Guangxi, W.T.Tsang 22699 (BM, A); Zhengkou, Jiangsu, C.P'ei 3215 (IBSC). CHINA SOUTH-CENTRAL:
Emei [Omei] Shan, Sichuan, W.P.Fang 14373 (IBSC); Chungtien [Zhongdian] Plateau, Yunnan, G.Forrest
12618 (E, BM). 38. TAIWAN: Ilan Hsien, C.C.Liao & D.S.Hsu 313 (A).
In China, India, Nepal, Thailand and Vietnam, the leaves are used medicinally, and the
shoots are eaten as a vegetable (E.F.Anderson, Econ. Bot. 40(4): 442–450, 1986; K.R.Kuebel
& A.O.Tucker, Econ. Bot. 42(3): 413–419, 1998; A.K.Gangwar & P.S.Ramakrishnan,
Econ. Bot. 44(1): 94–105, 1990; N.P.Manandhar, Econ. Bot. 49(4): 371–379, 1995). Forms
with variegated leaves and double flowers are grown as ornamentals in gardens in Europe
and the U.S.
9
SPECIES PLANTARUM —FLORA OF THE WORLD
MAPS
Number in brackets refers to the page on which the taxon is described.
10
2005
MAPS
1. Saururus cernuus (2)
2. Saururus chinensis (4)
3. Gymnotheca chinensis (5)
4. Gymnotheca involucrata (6)
5. Anemopsis californica (7)
6. Houttuynia cordata (7)
11
INDEX
Accepted names are in roman, synonyms and doubtful names in italic.
Principal page references are in bold, figures and plates in italic.
Anemia Nutt. 6
californica Nutt. 6
intermedia Copel. ex M.E.Jones 6
Anemopsis Hook. & Arn. 1, 6
bolanderi C.DC. 6
californica (Nutt.) Hook. & Arn. 6, 11
var. typica Kelso 6
var. subglabra Kelso 6
ludovici-salvatoris Willk. 6
Aponogeton
natans (L.) Engl. & K.Krause 5
Circaeocarpus 1
Gymnotheca Decne. 1, 5
chinensis Decne. 5, 8, 11
involucrata 5, 6, 11
weinmannifolia hort. ex L.Gentil 5
Houttuynia Thunb. 1, 7
bolanderi (C.DC.) Benth. & Hook.f. 6
californica (Nutt.) Benth. & Hook.f. 6
cordata Thunb. 7, 8, 11
f. polypetaloidea T.Yamaz. 9
f. viridis J.Ohara 9
foetida hort. ex Loudon 7
Mattuschkia J.F.Gmel. 2
aquatica (Walter) J.F.Gmel. 2
Polypara Lour. 7
cochinchinensis Lour. 7, 9
cordata (Thunb.) Kuntze 7
Saururaceae 1
Saururopsis Turcz. 2
chinensis (Lour.) Turcz. 2, 4
cumingii C.DC. 4
var. japonica C.DC. 4
Saururus L. 1, 2
aquaticus Walter 2
cavaleriei H.Lév. 5
cernuus L. 2, 11
f. submersus Glück 2
cernuus auct. non L. 4
chinensis (Lour.) Baill. 3, 4, 11
loureiri Decne. 4
lucidus Donn 2
natans L. 5
sinensis Teijsm. & Binn. 4
Spathium Lour. 2
chinense Lour. 2, 4
12