OVERVIEW OF LIRIOPE AND OPHIOPOGON (RUSCACEAE)
NATURALIZED AND COMMONLY CULTIVATED IN THE USA

Guy L. Nesom

2925 Hartwood Drive

Fort Worth, TX 76109, USA

www.guynesom.com

ABSTRACT

The Asian natives Liriope spicata, L. muscari, Ophiopogon jaburan, and O. japonicus have
been reported to be naturalized at various localities in the USA; rq>orts of O. jaburan have been
based on mis identifications. The naturalized occurrence of two Liriope species in Texas is
documented here: (1) L. spicata occurs abundantly in large colonies along a forested creek within a
city park in Fort Worth (Tarrant Co.) and it has been collected from a bottomland hardwood forest in
the Big Thicket National Preserve; (2) a large plant of L. graminifolia is growing in low woods in a
city park in Euiess (Tarrant Co.). A key, brief descriptions, and basic synonymy are provided for the
naturalized species as well as for L. gigantea. O. intermedins. O. jaburan, and O. planiscapus, which
also are sold and commonly cultivated in the USA. An provides representative

photos of the species in cultivation and in naturalized habitats. Liriope gigantea sensu stricto
(colonial from stolons) differs in habit from the species now generally sold as L. gigantea (caespitose)
— no species identified as L. gigantea has been recognized to occur natively in Asia. The name
Liriope exiliflora is treated as a synonym of L. muscari: plants abundantly cultivated in the USA and
often identified as L. exiliflora are instead L. graminifolia.
KEY WORDS: Liriope, Ophiopogon, cultivars, Ruscaceae, Liliaceae, naturalized, Texas, USA

Species of Liriope Lour, and Ophiopogon Ker Gawl. are commonly and abundantly planted
in the southeastern and southcentral USA in various ways in landscapes, particularly as ground cover
and border plants. All are characterized by basally tufted, linear, evergreen leaves and often are
known as "lilyturf," "mondo grass," "Aztec grass," or "border grass," or sometimes as "monkey
grass." At least one of them has been called "table grass," alluding to its graceful appearance in a pot
placed on a table or desk As a group, species of both genera are sometimes referred to as
"liriopogons," a termed introduced by Skinner (1971), generally supplanting the earlier "liriomondos"
suggested by Bailey (1929).

Four species of liriopogons have been reported as naturalized in the LISA: Liriope mmcari
(Dene.) L.H. Bailey, L. spicata Lour., Ophiopogon jaburan (Sieb.) Lodd., and O. japonicus (Thunb.)
Ker Gawl. (USDA, NRCS 2010; Kartesz 2010). Liriope graminifolia (L.) Baker' is added to this list
in the current report and O. jaburan is removed, as reports of the latter have been based on
mis identifications. Of these species, apparently only O. japonicus has been reported to grow outside
of cultivation anywhere else in the world — from one locality in Brazil (GCW 2010; Institute Horus
20 10). For the treatment of Liliaceae in Flora of North America North of Mexico (FNANM), Utech
(2002) noted that Liriope muscari, L. spicata, and Ophiopogon jaburan "have been reported as
escaped in the flora . . . but "are not clearly naturalized" and he did not provide a treatment for either
of the genera.

In addition to the species reported as naturalized in the USA, the present account includes

Liriope gigantea H. Hume, Ophiopogon intermedins D. Don, and O. planiscapus Nakai, which also
are sold and cultivated in the southeastern USA. An overview by Hume (1961) of liriopogons
cultivated in the USA included O. "arabicus" as distinct, but this is a horticultural name for a cultivar

Nesom: Uriopeand Ophiopogomn the USA

of O. planiscapus. Hume and Morrison (1967) included O. jaburan, O. japonicus, O. planiscapus, L.
exiliflora (here regarded as a synonym of L. muscari), L. gigantea, L. graminifolia (tenuously
distinguished by Hume and Morrison from L. exiliflora), L. muscari, and L. spicata in their account of
commonly cultivated lilyturfs. Broussard (2007) studied 19 cultivars of Liriope and Ophiopogon and
identified seven species among them — all of them are included in the present manuscript. A brief
note on the remaining three or four species of Liriope, to complete the genus, also is included.

Fantz (1993) observed that "names of available liriopogons cultivated in the United States
based on literature and catalogues [total] more than 20 named species and more than 100 named
cultivars, even after elimination of similar names" — he listed 26 species. In a follow-up publication
(Fantz 1994), he listed 22 species and 88 cultivars and noted (Fantz 1995) that "a minimum of
fourteen morphologically defined species are in cultivation in the United States" — 6 of Liriope and 8
of Ophiopogon. In his last two most recent publications, Fantz has provided a key and descriptions
for 6 species of Liriope (2008b) and for 14 of Ophiopogon (2009), documenting all as cultivated in
the southeastern USA or at least for sale in the regional horticultural trade. Fantz's studies (at North
Carolina State University in Raleigh) were supported by observations on plants of many or most of
the species and horticultural forms that he cultivated over 15 years.

Species of Liriope and Ophiopogon are native to southeastern Asia and the most
comprehensive accounts of both genera are in the Flora of China (Chen & Tamura 2000a, 2000b) and
the Flora of Japan (Tamura in press). Studies of species groups have been published by Hasegawa
(1968), Hara (1984), Tamura (1990), Yang and Li (1990a, 1990b), Yamashita and Tamura (2001),
and Tanaka (1998-2000, 2001). Except for the recent reports by Fantz. most accounts of liriopogons
in the USA have included only a few species (e.g., Bailey Hortorium Staff 1976; Walters et al. 1984;
Adams 1989; Deputy 1999). Regarding potentially naturalized species, beyond a specimen citation or
simply the inclusion of the listed name in a floristic account, there appears to be little or no
in any of these in North American taxonomic literature.

The present report documents Texas occurrences of Liriope spicata and L. graminifolia
outside of cultivation and provides a taxonomic and photographic overview of commonly cultivated
liriopogon species in order to provide a readily available source for their identification in the USA A
detailed account of liriopogons naturalized in Alabama is provided by Spauiding et al. (2010). For
critical studies botanist-- should have primary Asian literature, especially from the Flora of China and
the Flora of Japan, and the recent assessments by Fantz. The treatment here of Liriope is parallel to
that of Fantz (2008b) except for the name of one species, but Fantz (2009) has treated 14 cultivated
species of Ophiopogon compared to the four included here. The article by Hume and Morrison
(1967) also is useful for its descriptions of common cultivars.

Methods.

For the present review, I have seen many plantings of Liriope gigantea. L. graminifolia, L.
muscari, L. spicata, Ophiopogon japonicus, O. intermedins, and O. planiscapus. The account here of
O. jaburan is from published sources. Unless otherwise specified, state records given here for
putatively naturalized populations are from summaries by Kartesz (2010) and the PLANTS Database
(USDANRCS2010).

internet, nurseries), I have attempted to align names of
common cultivars with species, but this should be taken only as a best approximation. Some cultivar
names apparently have been duplicated (e.g., a "Variegata" has been developed in more than one
species) and inconsistencies are widespread in identification of species and cultivars (see details and
examples by Fantz 1993, 1994), even as distributed by growers. Even patent applications for new
cultivars are apparently not required to document the parentage (e.g., see applications shown at

Nesom: Uriopeand Ophiopogomn the USA

internet sites "PatentStorm," "PatentGenius," "FreePatentsOnline," etc. — or search the US Patent
Office Collection <http://patft.uspto.gov/netahtml/PTO/search-adv.htm>).

Synonymies of scientific names are derived from primary literature and floristie treatments of
Chinese and Japanese species (as cited below). The provenance is indicated for types of all of the
names included here and 1 have examined types where photographs or digital images are available (as
indicated in the typification summaries; images via website of individual herbaria and JSTOR). Hi-
res images of the holotype of Convallaria spicata Thunb. were made available by staff at UPS and
those photos are included in the manuscript as Figures 1 and 2.

A set of phulo-, taken mo-th b\ the authui i^ included in thw piescnl report (see
Appendix). Photos identified by cultivar name can be found on a number of commercial internet
sites, e.g., "acemondo.co.nz," "davesgarden.com," "gogardennow," "groundcovertogo.com," and
"liriope.com." A good site for a diversity of high-quality photos of Liriope and Ophiopogon, with
interesting notes by J.C. Raulston, is at <www.cultivar.org>. The dissertation by Broussard (2007;
online) also has a series of good photos of cultivars in situ and of representative herbarium specimens.

Liriope and Ophiopogon at generic rank.

Liriope includes about eight species distributed in Pakistan, China (including Taiwan), Korea,
and Japan to Indochina (Vietnam, Laos, Cambodia) and the Philippines. Ophiopogon includes about
54 species (fide Conran & Tamura 1998) or 65 species (fide Chen & Tamura 2000b) in essentially the
same area as Liriope but definitely known from other Himalayan countries and most of Indochina
(see distribution of O. intermedins, below) as well as Malesia. The two genera are closely related and
vegetatively similar — the most significant differences are in floral morphology (see key below).
Both are now placed in the Ruscaceae (Rudall et al. 2000; Nyffeler & Eggli 20 10).

A dendrogram of 18 popular cultivars studied by Mcharo et al. (2003) — based on AFLP
markers — diverges somewhat from expected groupings according to the putative name assignments
here, which lead the authors to suspect that Liriope and Ophiopogon might be conspecific.
Differences in floral morphology, however, appear to be consistent and floristie treatments and other
Asian studies have not found placement of species to genus problematic.

The genera Liriope, Ophiopogon, and Peliosanthes Andrews (ca. 11 species, south China to
Indochina, Malesia, and India; or 1 species, fide Jessop 1976; also see Yamashita et al. 2002) have
been considered to comprise the tribe Ophiopogoneae Endl. (e.g., Cutler 1992; Conran & Tamura
1998) or even a separate family, the Ophiopogonaceae Meisn. (Conran 1989), defined by tufted
JWus r j ss likw lents i ^piieli 1 *. infkiescOKe me^ulm md ^i t h < nis ent, 1-seeded fruits,
and sarcotestal seeds. It is clear that Liriope and Ophiopogon are sister taxa, but the relationship of
Peliosanthes has been shown to lie elsewhere (Rudall et al. 2000), reducing the Ophiopogoneae to
only the two genera.

Notes on species biology.

a. Chromosome number.

At least some of the widespread species of Liriope and Ophiopogon are variable in ploidy
level (e.g., see L. graminifolia, L. muscari, L. spicata, O. intermedins, O. japonicus; chromosome
number summaries at the end of morphological descriptions are from Zhang 1998). This at least
suggests that the morphological variation may be partitioned to a greater extent than currently
recognized. If alloploidy has been involved in the formation of supra-diploid levels, attempts to
understand relationships among species may be minimally meaningful without a detailed knowledge
of genomic constitution of the plants under investigation. Chromosome number variation appears to

Nesom: Liriope and Ophiopogomn the USA

have originated in natural populations; I have not encountered any discussion or indication of
hybridization involved in the production of cultivars.

Fukai et al. (2008) estimated ploidy levels (using flow cytometry) of cultivars they identified
among six species of Liriope and Ophiopogon: L. minor (2x), L. spicata (6x), O. jaburan (2x), O,
japonicus (4x), O. planiscapus (2x), and O. platyphylla (4x). Apart from tliis report, I have not
encountered information regarding ploidy levels of cultivars.

h. Flowers.

The occurrence of inferior to semi-inferior ovaries in Ophiopogon is viewed by Rudall (2002)
as one among numerous multiple and independent origins of epigyny within monocots in general and
Asparagales in particular, probably linked with the evolution of different pollination syndromes. As
in Liriope, flowers of most taxa of Asparagales are hypogynous, the primitive condition; the
predominantly epigynous to perigynous flowers of Ophiopogon are specialized. Zhang (1998) has
noted, however, that the position of the ovary is variable in both Liriope and Ophiopogon, sometimes
even within a single species, and thus is not a consistently diagnostic feature.

Pedicels of these plants in early development are similar in color and texture to the corolla
and floral tube (in Ophiopogon) and corolla (in Liriope). In Liriope, where a floral tube is essentially
absent, the "pedicel" lias been described in some literature as articulated distally, i.e, directly beneath
the corolla. In Ophiopogon, the "pedicel" has been described as articulated medially, indicating the
point of division between the slender floral tube (the "perigone," iide Fantz 2008a), and the actual
pedicel. During fruit development or after abscission of the corolla, the persistent pedicel in both
genera becomes more clearly differentiated, appearing stramineous and somewhat sclerifled. At least
in O. Jaburan and O. japonicus, and presumably other species of Ophiopogon with semi-inferior
ovaries, the pedicel and perigone may remain connected and persistent as a unit subtending the fruit
until after it is released.

In some of cultivars of Liriope, flowers apparently never open and eventually are deciduous
still in the form of buds. Hume and Morrison (1967) specifically noted that this is true of L. muscari
"Christmas Tree." The observation by Hume (1961) and Hume and Morrison (1967) — and
corroborated here at least for some cultivars of L. muscari and L. graminifolia — that the sequence of
flower development is irregular over the inflorescence was questioned by Zhang (1998), who noted
that opening of flowers is generally acropetal. An acropetal sequence can be seen in some young
inflorescences (see photos of L. muscari in the Online Appendix).

c. Stolons and roots.

Some of the species in the present account reproduce clonally by shallow, laterally spreading
stolons. These are slender and whitish underground stems with narrowly triangular scale leaves at the
relatively widely spaced nodes, usually unbranched, sometimes producing adventitious roots, and
producing a new leafy plant at each stolon apex. One or more stolons may be produced from a single
leafy plant. Fantz (2008a) termed these structures rhizomes, restricting the use of stolon to creeping,
above-ground stems. In the liriopogons, these might be termed stolon-like rhizomes, but they are
closest to what are generally identified as stolons in plants over a wide range of taxonomically diverse
species (see excellent summary and discussion of stolons and rhizomes in Wikipedia). Stolons and
rhizomes are not completely dichotomous in distinction, and the term rhizome might be appropriately
applied to very short stolons sometimes produced in Liriope muscari (see comments below).
Representative stolons are shown in the Online Appendix.

Elongating stolons usually are not formed in Liriope muscari and Ophiopogon jaburan,
which are "caespitose" forms that increase and slowly spread by production of new shoots from

Nesom: Uriopeand Ophiopogomn the USA

lateral buds in the caudex crown. Hume and Morrison (1967) noted that "an old clump of L. muscari
eighteen inches in diameter may give as many as one hundred and fifty separate pieces'' ("bibs,'"
daughter plants with few leaves, fide Fantz 2008a), presumably from crown buds, but the manner of
reproduction was not specified.

Some forms of cultivated Liriope muscari, however, do produce short stolons (see comments
below under L. muscari and photos in the Online Appendix), and stolon production also appears to be
variable in L. gigantea (see comments under L. gigantea). The same may be true in other primarily
caespitose, "clumping" species (e.g., perhaps cuitivars of Ophiopogon intermedins), Fantz (2008a, p.
335) noted that "weakly rhizomatous plants" [of liriopogons] have short stolons, "with daughter
plants borne close to the mother plant and their roots intertwining. This growth pattern will form a
dense carpet or mound over the ground with age. with original planting sites merging together,
providing an attractive ornamental appearance."

Liriopogons are generally described j . "clumping oi "spreading" in horticultural
information but below-ground parts are almost never described. Unequivocally "spreading" or
"colonial" forms, which sometimes are also termed "invasive," are L. spicaia. L. graminifolia, and
Ophiopogon japonicus. Caespitose or primarily caespitose forms are used for borders and for
plantings where a certain persistent geometry is desired.

Most of the liriopogon species in cultivation in the USA produce tuber-like ellipsoid
swellings, sometimes in tandem, usually at or near the distal ends of the fibrous root system (photos
in Online Appendix). Steroidal saponins, steroidal glycosides, and sesquiterpene glycosides have
been isolated from tuberous swellings of Liriope muscari, L. spicata, O. japonicus, and others (e.g.,
Ahmad & Basha 2006; Cheng et al. 2004, 2006; Yu et al. 1996). The root "tubers" of L. spicata are
used medicinally in China and the species is widely cultivated there for that reason. These organs
have not been observed to have a rq^roductive function.

KEY TO THE COMMONLY CULTIVATED SPECIES

1. Flowers erect, terminal at the tip of a strict pedicel, corollas cupulate to rotate; ovaries superior
(flowers hypogynous); anthers free, oblong in outline, apically obtuse, dehiscing apically and
appearing poricidal. filaments longer than or as long as anthers; stigmas entire; fruits blackish

LIRIOPE

2. Plants caespitose, without stolons or weakly stoloniferous or rhizomatous; leaves (5-)8-16(-23)
mm wide, (15-)30-70 cm long.

3. Leaves mostly 24-55 cm long, relatively thin, flexuous and arching 45° to 140° or more;

scapes usually equal or longer than the leaves, sometimes shorter 1. Liriope muscari

3. Leaves mostly 50-80 cm long, relatively thick, usually relatively stiff, slightly arching 45" to
90°; scapes usually distinctly shorter than the leaves 3. Liriope gigantea

2. Plants colonial, from slender stolons; leaves (2-)3-8(-10) mm wide, (14-)18-60 cm long.

4. Scapes (15-)20-44(-53) cm long, mostly equal or longer than the leaves and the
inflorescences conspicuous; corollas purple to lavender; inflorescence rachis (4-)5-13 cm long,

(20-)45-160-flowered 2. Liriope graminifolia

4. Scapes 12-29 cm long, mostly much shorter than the leaves and the inflorescences mostly
obscured; corollas mostly white; inflorescence rachis 2-5(-8) cm long, (20-)45-72 -flowered
4. Liriope spicata

Nesom: Uriopeand Ophiopogomn the USA

1. Flowers nodding, on a recurving pedicei/floral tube, corollas mostly campanulate; ovaries inferior
to semi-inferior (flowers epigynous to perigynous); anthers more or less connate in a column,
narrowly lanceolate in outline, apically narrowly attenuate-acute, dehiscing longitudinally, filaments
much shorter than the anthers (anthers subsessile); stigmas 3-lobed; fruits commonly blue, sometimes
blackish OPHIOPOGON

5. Leaves 2-4 mm wide; scapes 6-12 cm long, much shorter than the leaves: inflorescence rachis

2-5 cm long, 5-10+-flowered; plants colonial 1. Ophiopogon japonicus

5. Leaves 4-15 mm wide; scapes (12-)15-50 cm long, shorter or longer than the leaves;
inflorescence rachis (2.5-)4-16 cm long, lQ-32-flowered or 45-250-flowered; plants colonial to
weakly colonial.

6. Leaves blackish purple to black in popular horticultural forms, 4— 9(-12) mm wide, (10-)25-

36 cm long; 10-32-flowered; plants strongly colonial 2. Ophiopogon planiscapus

6. Leaves green or variegated, 7-15 mm wide, (20)25-60 cm long; 45-250 flowered; plants
weakly colonial.

7. Rachis 6-10.5 cm long, fascicles 12-20; flowers 3-7 per fascicle calyx tube (perigone) 6-
8 mm long; pedicels (in flower) 9-1 1 mm long; leaves green or variegated

3. Ophiopogon jaburan

7. Rachis (8-)17-21 cm long, fascicles (30-)45-55; flowers (l-)3-6 per fascicle; calyx tube
(perigone) 1-2 mm long; pedicels (in flower) 2-5 mm long; leaves variegated
4. Ophiopogon intermedius

1. LIRIOPE MUSCARI (Dene.) L.H. Bailey, Gentes Herb. 2: 35. 1929. Ophiopogon muscari
Dene., Fl. Serres Jard. Eur. 17: 181. 1867-1868. TYPE: China ("Habitat in insula Hong
Kong?").

Convallaria spicata Thunb. in Murray, Syst. Veg., ed. 14, 334. 1784 [also in Fl. Jap. 141. 1784].
Ophiopogon spicatus (Thunb.?) Ker Gawl,, Bot. Reg. 7: pi. 593. 1821 [nom. invalid.] [non
Hook 1862]. Liriope muscari (Decne.) L.H. Bailey var. exiliflora L.H. Bailey [nom. nov.],
Gentes Herb. 2: 37. 1929. Liriope exiliflora (L.H. Bailey) H. Hume, Baileya 9: 144. 1961.
Liriope muscari forma exiliflora (L.H. Bailey) H. Hara, J. Jap. Bot. 59: 37. 1984. TYPE:
Japan. "Crescit prope Nagasaki" (fide Flora Japonica, p. 141), C.P. Thunberg s.n. (holotype:
UPS digital image!. Figs. 1 and 2 of present account, photograph in Bailey 1929 as Fig. 7);
probable isotypes: BM digital images!, the plants represented only by scapes).

Wang and Tang (1951) proposed the combination Liriope platyphylla var. exiliflora
(Bailey) Wang & Tang, but they did not provide the basionym and the name is invalid. See
further comments below, under "The status of the name Liriope exiliflora."

Liriope graminifolia (I,.) Bakes' var. densifolia Maxim, ex Baker, J. Linn. Soc, Bot. 17: 500. 1879.
Liriope spicata (Thunb.) Lour, var. densifolia (Maxim, ex Baker) C.H. Wright, J. Linn. Soc,
Bot. 36: 79. 1903. TYPE: "Japonia, Formosa, China borealis, et insulae Loo Choo." As
synonym fide Chen and Tamura (2000b), Tamura (in press).

Ophiopogon spicatus (Thunb.) Ker Gawl. var. communis Maxim., Bull. Acad. Imp. Sci. Saint-
Petersbourg 15: 85. 1871. Liriope muscari var. communis (Maxim.) Nakai. Bot. Mag.
(Tokyo) 43: 776. 1934, Liriope muscari var. communis (Maxim.) P.S. Hsu & L.C. Li, Acta
Phytotax. Sin. 19: 460. 1981. TYPE: Japan. Possible isotype: Prov. Senano et Nambu, 1864,
1865, and 1866, S. Tschonoski s.n. (NY digital image!); a probable duplicate of this, but
giving the date as 1864 and the collection number as Tschonoski 1874. is at P, digital image!.
Basal parts not collected. As synonym fide Chen and Tamura (2000b), Tamura (in press).

Liriope yingdeensis RH. Mao, Acta Sci. Nat. Univ. Sunyatseni 1982: 75. 1982. TYPE: China,
Guangdong Province. As synonym fide Chen and Tamura (2000b), Tamura (in press).

Nesom: Uriopeand Ophiopogomn the USA

Stolons usually absent, short if present. Roots with tuberous swellings but usually few,
absent. Leaves (15-)24-55(-65) cm long, 6-12(-23) mm wide, stiff and erect to flexuous
and arching, (5-)12-25-nerved, margins minutely serrate distally, apices acute to obtuse or rounded.
Scapes erect, 24-60(-70) cm long, at least some distinctly longer than the leaves, others often shorter,
sometimes all about the same height as the leaves, subterete to slightly flattened, especially distally,
or distinctly flattened proximally and distally, green to purple. Inflorescence (50-)100-220(-300)
flowered, rachis 6-20 cm long, narrowly columnar to broadly cylindric with loosely to densely
arranged fascicles; bracts lanceolate to triangular, proximal 3-6 mm long. Flowers mostly in
fascicles of (1 or 2-)3-?, usually reduced to (l-)2-5 distally; pedicels (2-)3-5 mm long; tepals
purple, lilac-purple, or lavender, elliptic-oblong to oblong, 3-4 mm long; floral tube 0.2-0.6 mm
long, less commonly essentially absent. Fruits globose, 4-8(-9) mm in diam., blue-black to dark or
blackish purple to nearly black at maturity. Seeds blackish purple at maturity, subglobose, 4-7 mm in
diam. 2n = 36, 72, 108, 112.

Native to China, Taiwan, Japan (including Ryukyu Islands). Native habitats include forests,
bamboo forests, scrub, shady and moist places in ravines and on slopes; 100-1 400(-2000) meters
elevation. Common names: BIG BLUE LIRIOPE, BLUE LILYTURF, BLUE-FLOWERED SNAKESBEARD
(fide Ker Gawler 1821). Representative cultivars: Big Blue (later known as Riegel Big Blue and
Original Big Blue), Bigun, Blue Spire, Border Gem, California Hybrid, Christmas Tree (sometimes
known Monroe No. 2), Cleopatra, C.T. Tanner, Curly Twist, DensifJora, Eleven-O-Three, Gold
Banded (v), Grandiflora, Green Midget, Hawk's Feather (v), fngwersen, Isabelle, Jeanette, John
Burch (v), Just Right, King Felix. Lilac Beauty. Majestic, Marant, Marc Anthony, Mayan Blue.
Moneymaker, Monroe White (sometimes known as Monroe No. 1), New Blue, New Wonder, Okina,
Paul Alden (v), Peedee Ingot, Purple Bouquet, Royal Purple, Silver Banded (v), Silver Dragon (v),
Silver Ribbon (v), Silvery Midget (v), Silvery Sunproof (v; see comment under L. graminifolia), Sno
Cone, Summer Beauty, Superba, Tidwell's Big Blue (now marketed as Tidwell's True Blue),
Traebert White, Variegata (v), Webster Wideleaf Fkwermg (Jun-)Jul-Aug. "(v)" = variegated.

Naturalized distribution in the USA Alabama, Georgia, Kansas, Louisiana (Lincoln Par.,

Thomas & Allen 1993), Maryland, Mississippi, South Carolina. A report of Liriope muscari from
Louisiana (MacRoberts et al. 2008); proved to be I ip cat i and it s an likch th.it ,t least some
others also are mis identifications of stoloniferous species. For example, images of 'infestations' of
monkeygrass identified as L. muscari (Forestry Images 2010; no indication of provenance except
"United States") probably are L. spicata. On the other hand, at least one Florida collection
(Wunderiin & Hansen 2008) identified as L. spicata apparently is L. muscari — Jackson Co.: Burks
1125, USF; the identification of Wilhelm 10159 —from Okaloosa Co., USF — as L. spicata probably
is correct. Diggs et al. (1999) noted that L. muscari in north central Texas is widely cultivated and
persists and spreads vegetatively in flower beds — the characterization of its vegetative spread
suggests that a stoloniferous species (e.g.. L. spicata or L. graminifolia vs. the primarily caespitose I.
muscari) was included in the concept.

Deputy 7 (1999) noted that Tidwell's Big Blue is a "spreading form ... not to be confused
with the clumping type Big Blue." Paul Fantz (pers. comm. 2010) also observed that two plants of
Tidwell's Big Blue, obtained from Tidwell's Nursery in 1987 and grown in research plots, proved to
be stoloniferous. Tidwell's Nursery (pers. comm. 2010, fide "Bo" Tidwell) affirms that at least
Tidwell's True Blue as now marketed is a 'clumping' form. See other comments on variability in
stolon production — under "Notes on species biology" and with photos in the Online Appendix. It
even seems a distinct possibility that some cultivars currently marketed as L. muscari may have been
derived from forms of L. graminifolia in which stolon production has been repressed or restrained.
Evidence for this might be sought through molecular studies.

Nesom: Uriopeand Ophiopogomn the USA

The wide variation in cultivars in habit and other features at least suggests that the genetic
poo! of Liriope muscari may be greater than in other liriopes. As noted by Fantz (2008b, p. 347),
"This is the only species with inflorescence variations that have lead to distinct cultivar selections/"
Consideration of L. platyphytta as a species distinct from L. muscari (see below) alsj anpha' izes ths.
taxonomic complexity of the lattter, and it seems likely that critical study may bring to light
additional taxa that can be justifiably recognized.

Status of the name Liriope exiliflora.

Bailey (1929) explicitly noted that his Liriope muscari var. exiliflora was intended as a
nomen novum (a replacement name at different rank) for Convallaria spicata Thunb. In Hume's
recognition of the taxon at species rank, the prior publication of Liriope spicata Lour. (1790) blocked
usage of Thunberg's epithet in the genus Liriope.

Chen and Tamura (2000a) interpreted the name Liriope spicata Lour, as a combination by
Loureiro based on C. spicata Thunb. As observed by Bailey (1929), however, Convallaria spicata
Thunb. (the type from Japan) and Liriope spicata Lour, (the type probably from China) are
heterotypic and represent different species.

The application of the name Liriope exiliflora to the plants commonly cultivated in the USA
stems from the interpretation by Flume (1961). From a photo of the LIPS type of Convallaria spicata
Thunb., which Bailey had renamed as L. muscari var. exiliflora, Hume was convinced that the plant
shows "a piece of rhizome attached to the base of the segment" and thus would be a match for the
colonial species that he saw in cultivation. Dr. Mats Hjertson, UPS Curator, has sent high resolution
images of the UPS specimen (Figs. 1 and 2in the present account) — the thickened axis from the base
interpreted by Hume as a rhizome is instead a cord-like fibrous root, probably contractile in function.
The type specimen of Convallaria spicata can be matched in aspect to many plants of L. muscari seen
in cultivation.

Reservations earlier expressed by Skinner (1971, p. 348) about the recognition of Liriope
exiliflora can now be seen as well-founded: "Its origin is obscure. H.H. Hume has accorded it species
rank, but a cultivar designation would have seemed more logical, pending assignment to a known
existing species."

In raising Liriope exiliflora to specific rank, Hume contrasted it in tabular form with L.
muscari. Another stoloniferous species, L. graminifolia was also included in Hume's overview, and a
comparison of his description of L. graminifolia -vs.- L. exiliflora shows this: leaves 35-40 cm long,
3-5 mm wide, 9-11-nerved -vs.- 'to 43 cm" long, 9-12 mm wide, 9-13-nerved; rachis 3.5-6,2 cm
long -vs.- 8-16.5 cm long; fascicles ?, mostly 3-flowered -vs.- 3-5, mostly 5-flowered. The leaf
width described by Hume for /,. exiliflora matches the leaves of the type specimen (= Convallaria
spicata, Thunb.) but is considerably broader than the dimensions recorded by Fantz (2008b) and those
here for the cultivated stoloniferous species identified as L. exiliflora. Thus even the identity of the
American plants regarded by Hume as L. exiliflora is not perfectly clear in his eoncqrt, and a distinct
species identified as L. exiliflora has never been recognized in Japan, even a half century after its
taxonomic uplift by Hume. As concluded here, the plants identified by Flume as L. exiliflora are
forms of L. graminifolia (see discussion below), the type of which is from somewhere in Asia outside
of Japan, since L. graminifolia does not occur in Japan.

In both the original and revised edition of his Manual of Cultivated Plants. Bailey included

only two species, Liriope spicata and L. muscari, treating var. exiliflora within the latter (Bailey
1924, 1949) as a form with leaves "often only 6 mm broad but sometimes nearly twice as broad."

Nesom: Uriopeand Ophiopogomn the USA

Both L. exiliflora and L. graminifolia ate listed in Hortus Third (Bailey Hortorium Staff 1976), the
latter noted as "probably not in cultivation" (perhaps just repeating earlier comments).

Placement of Liriope exiliflora as a synonym of L. muscari accounts for several previous
discrepancies: (1) the morphological discordance between the type specimen of Convallaria spicata
Thunb. and plants identified as L. exiliflora in the American horticultural trade, (2) the absence of
plants identified as L. graminifolia in the American horticultural trade, and (3) the absence of a
distinctive species identified as L. exiliflora in Japan, where the type specimen was collected. For the
Flora of Japan, Tamura (in press) has treated Liriope muscari var. exiliflora L.H. Bailey as a synonym
of L. muscari sensu stricto, and the present study is in agreement.

Status of Liriope platyphyUa.

Liriope platyphyUa was proposed as a new species in 1951 and the name has engendered
taxonomic confusion, especially after being used by Ohwi (1953) for all of the L. muscari-Vks plants
of Japan. Recent Asian floras (Chen & Tamura 2000a; Tamura in press) have placed L. platyphyUa in
synonymy under L. muscari, but it has been maintained as a species distinct from L. muscari by Fantz
(2008b), who observed that "Plants bearing the name L. platyphyUa originating from China or
European nurseries (who obtained them from China) are very distinct from cultivated selections of
U.S. plants known as L. muscarf (p. 345). And (p. 348) "The inflorescences overtop the leaves
much higher than L. muscari, bearing more fascicles of flowers than any other species. However,
fruiting pedicels will bear dried, persistent perianth members, which is less desirable as an ornamental
trait when fruit typically abort. Rabbits continually ate the leaves of this species in research beds,
ignoring L. muscari and other species, which exhibited infrequent feeding."

Liriope platyphyUa F.T. Wang & T. Tang, Acta Phytotax. Sin. 1: 332. 1951. SYNTYPES: China:
Anhwei, T.N. Liou &. P.C. Tsoong 2077; Kiangsu, Y.L. Keng 2749 — cited by Wang and
Tang as "Type specimens." The herbarium of deposition was not specified for either, but the
authors were working from "Academia Sinica" and presumably the specimens are at PE in
Beijing. Fantz (2008b) noted that he had not been able to obtain type material for study

Liriope spicata var. latifolia Franch, PI. David. 296. 1884. TYPE: China. Kiangsi, Kiukiang
Mountains, A. David 854 (P). As synonym fide Chen and Tamura (2000b), Tamura (in

Liriope muscari forma latifolia (Makino) H. Hara, J. Jap. Bot. 59: 37. 1984. Liriope graminifolia var.

latifolia Makino, Yagaihakubutsu 3: 165. 1933. TYPE: Japan.

In synonymy of Liriope platyphyUa, Wang and Tang cited "Liriope muscari Bailey [1929]
non Ophiopogon muscari Decne. [1868]," but Bailey's name as a new combination was clearly based
on Decaisne's basionym and thus is homotypic with it. The authors described their new species as
"rhizomatosa," and it is possible that they conceived of it as distinct in growth habit from "Liriope
muscari sensu Bailey." They also cited in synonymy "Ophiopogon spicatus Hook. [1862] non
Liriope spicata Lour." (thus referring to Convallaria spicata Thunb. = Liriope exiliflora, see
comments below — which is stoloniferous). Even if they were proposing L. platyphyUa as a
replacement name for Convallaria spicata, Bailey had already provided that in 1929 as L. exiliflora.
In any case, citation of L. muscari, a valid species heterotypic with/.. platyphyUa, in synonymy oft.
platyphyUa appears to render' the latter invalid. (617) 495-0794

Among collections from the Harvard Herbaria, I have seen duplicates of several cited by
Wang and Tang as paratypes of Liriope platyphyUa: [China]. Chekiang, Cheo & Wilson 2805 (GH);
Hunan, Fan & Li 565 (GH); Kweichow, Steward, Chiao, & Cheo 77 (A, GH). From these, and from
the illustration in Wang and Tang (1978), the description in the protologue, and the implication of the
epithet, it appears that L. platyphyUa refers a relatively broad-leaved variant that is treated by current
floristicians in China and Japan as a synonym of L. muscari — representative collections of this form

Nesom: Uriopeand Ophiopogomn the USA

are shown in Figs. 5 and 6, compared to collections of typical L. muscari in Figs. 3 and 4. I have seen
collections generally matching the L. platyphytta variant from Japan and Taiwan and a cluster of
provinces of southeastern China (Jiangsu, Anhui, Zhejiang, Jiangxi, Hunan, and Guangxi). Typical L.
muscari appears to be restricted to eastern provinces of China (Shandong, Jiangsu, Anhui, Jiangxi,
Zhejiang, Jiangxi, Fujian, Guangdong, and Hainan) and it also occurs in Japan and Taiwan. 1 am not
able to adequately assess whether or not intergradation indicates that L. platyphylla and L. muscari
are conspecrfic.

The original description of Liriope /. h i \ I noted th it it is hi/om itous Dut m ie ot tju.
plants I have seen that otherwise match L. platyphylla, either cultivated or from the native range, have
been rhizomatous. Fantz (2008b) characterized the species as caespitose and forming tufted clumps.

Some variants in the USA horticultural trade match the leaf morphology of Liriope
platyphylla. Fantz's key (2008b) key does not directly compare L. platyphylla and L. muscari, but the
couplet below is derived from his key and descriptions.

1. Scapes 55-60 cm long; rachis 18-24 cm long, crowded, ca. 3/4-4/5 as long as the peduncle;
fascicles 60-80; perianth tube prominent, 1-1.2 mm long; corollas lavender; anthers 1.6-1.8 mm
long; outer bracts obovate-elliptic, hyaline wings to near apex Liriope platyphylla

1. Scapes 20-30 cm long; rachis 6-12 cm long, loose, ca. 1/3 as long as the peduncle; fascicles 35-
50; perianth tube inconspicuous, 0.1-0.4 mm long; corollas violet to purple or white; anthers 1.5 mm
long: outer bracts linear with abrupt, ovate, hyaline wings basally Liriope muscari

It may be possible to recognize another broad-leaved variant of Liriope muscari sensu lato.
Plants with exceptionally long, thick leaves and long scapes (see representatives in Figs. 7 and 8)
occur in provinces mostly west of the "platyphylla" variants (Sichuan, Yunnan, and Guizhou, and
overlapping or parapatric with "platyphylla" in Hunan). The leaves are relatively even in width rather
than prominently broadening distally as in the "platyphylla'" form. More detailed study would be
necessary to accurately portray the variation.

2. LIRIOPE GRAMINIFOLIA (L.) Baker, J. Linn. Soc, Bot. 14: 538. 1875. Asparagus

graminifolius L., Sp. PI., ed. 2, 1: 450. 1762. Dracaena graminifolia (L.) L., Syst. Nat. (ed.
12) 2: 246. 1767. Mondo graminifolium (L.) Koidz., Bot. Mag. (Tokyo) 40: 333. 1926.
Ophiopogon graminifolius (L.) H.R. Wehrh., Gartenstauden 1: 187. 1929. TYPE: "Habitat in
Asia."

A type has not been designated for Asparagus graminifolia (fide Jarvis 2007), but
original material is in herbarium LINN (Savage No. 435.5 digital image!;
<http://www.linnean-online.org/5244/>). The plant is in flower, clearly Liriope, and lias narrow
leaves (2-2.5 mm wide) and 11-25 cm long. The inflorescence overtops the leaves — the
scape (peduncle plus rachis) is 27 cm long and the rachis is 11 cm long. The fascicles are
well-separated and each is composed of (l-)2-4 flowers with ovate floral bracts; the flowers
total about 85. The basal parts of the plant, critical in its identification, were not collected but
the narrowly linear leaves and overtopping inflorescence give some confidence that it
represents the species currently identified in Asia as L. graminifolia.
Liriope crassiuscuia Ohwi, Acta Phytotax. Geobot. 12: 108. 1943. TYPE: China. As synonym fide
Chen and Tamura (2008a).

Stolons creeping, slender (fide protologue and observation). Roots with tuberous swellings.
Leaves (14-)18-54 cm long, (2-)3-8(-10, -12 fide Fantz) mm wide, flexuous and arching, (6-)8-15-
nerved, margins minutely serrulate distally, apices acute to obtuse to rounded. Scapes erect, (15-)20-
44(-53) cm long, usually longer than the leaves, sometimes subequal to slightly shorter, subterete to

Nesom: Liriope and Ophiopogomn the USA

strongly flattened, especially distally. Inflorescence (20-)45-160-flowered, rachis (4— )5— 13 cm
long, narrowly columnar with distinctly separated fascicles; bracts ovate, proximal 4-5 mm long.
Flowers in fascicles of (l-)2-7, usually 1-3 distally; pedicels 2-5 mm long; tepals light purple to
lavender, oblong to oblong-elliptic, 3-4 mm long; floral tube 0.5-1 mm long. Fruits globose, 4-5
mm in diam., blackish at maturity. 2n = 72, 108.

Native to Pakistan and northwestern Himalayas. Vietnam, and China (incl. Taiwan). Native
habitats include forests, thickets, shady places along ravines, grassy and rocky places; ca. 10-2300
meters elevation. Common names: GRASS LILYTURF, CREEPING LILYTURF. Representative
cultivars: Angustifolia \tiaku-iiir4iigt »\ =Silvery Sunproof?), Samantha ("pink liriope"). Some
commercial websites equate the names Ariaka-janshige and Silvery Sunproof; a few describe it as
"spreading" but photos mostly show caespitose plants. Flowering (Jul-)Aug-Sep.

Reported here for the first time (see below) as outside of cultivation in the USA. Hume
(1961, p. 147) noted that "In the Lower South it [Liriope exiliflora sensu Hume = L. graminifolia] is
commoner than any other species of Liriope." Fantz (2008) also noted that it is very common in the
landscape, and the present study confirms its abundance in cultivation, at least in Texas, Louisiana,
and Arkansas.

In a taxonomic account of the Asparagaceae, Baker (1875) disposed of various names outside
of his concept of Dracaena. His entry for the liriope was this: "D. GRAMINIFOLIA, L. = Liriope
Sdmimloli ' Bake? = Ophiopogon spicatus, Gawl." Fantz (2008b) treated Liriope graminifolia as a
synonym of L. spicata Lour., acceding to a precedent that he cited from the 2007 International Plant
Names Index (IPNI), but Baker's citation of "Ophiopogon spicatus Gawl." would have been in
reference to the basionym Convallaria spicata Thunb., which is regarded here as a synonym of L.

Examination of the types of Liriope graminifolia (L.) Baker and L. spicata Lour, (see
references above to the images) indicates tliat they are not the same species (and, of course, the
Linnaean name is the earlier). Liriope spicata is distinguished by its longer and narrower leaves,
shorter scapes (relative to the leaves), much shorter and fewer-flowered rachises, and white flowers.
The description of L. graminifolia formulated here from first-hand observations of cultivated plants in
the USA. is remarkably similar to the description of L. graminifolia in. the Flora of China (Chen. &
Tamura 2000a), presumably based on a wide geographic sample of native plants.

Liriope graminifolia and L. muscari, as identified here, are similar in aspect and can easily be
confused in identification, especially if the stolons of L. graminifolia are restrained by a barrier from
spreading — in both species the scapes characteristically overtop the leaves and the flowers are
purplish. The onh con-sisienl dt^gno^iic feature is the growth habit, stoloniferous in L. graminifolia,
caespitose in L. muscari. Fantz (2008, p. 346) described a useful technique for observing the
difference without digging: "A good field observation is to use your hands to separate the leaves in a
Liriope patch from above. It will be easy to observe patches of bare soil between bibs within the
clump of [L. graminifolia] because of its rhizomatous [stoloniferous] habit. The bibs will be close
together in caespitose L. muscari, making patches of bare soil more obscured or hidden."

Fantz also noted that "abundant fruit set also indicates Liriope [graminifolia], as fruit
commonly abort in L. muscari, leaving pedicellate racemes lacking or bearing very few fruit."

Nesom: Liriope and Ophiopogomn the USA

Liriope graminifolia naturalized in Texas.

The account here is the first report of Liriope graminifolia naturalized in the USA

Texas. Tarrant Co. : city of Euless, Bob Eden Park along S side of West Mid Cities Blvd., low woods
of Uhnus americana, Carya illinoiensis, Querent shumardii, Fraxinus pennsylvanica, and Moms
rubra, along slough immediately tributary to large creek, producing abundant fruit, 10 Oct 2010,
Nesom FW1 0-25 with Robert George (BRIT).

A single large plant with stoloniferous offsets was encountered (see photos in Online
Appendix). There is no obvious source for the species in the immediate vicinity but large amounts of
water move through the habitat in heavy rains and it is possible that a plant fragment was washed into
the site from a neighborhood outside the park. Liriope graminifolia was observed in cultivation in a
nearby neighborhood. Reproduction through seed is possible, especially in view of the abundant fruit
production — but then one might expect that escapes would be more commonly reported, since the
species is so commonly cultivated in the Southeast. Fruits of the isolated plant in Bob Eden Park
perhaps were developed apomictically.

3. LIRIOPE GIGANTEA H.H. Hume, Baileya 9: 148. 1961. TYPE: Presumably Asia, but
provenance otherwise unknown. "The source from which it came originally is obscure. It
was found growing in a small area near the house at Millpond Plantation, Thomasville,
Georgia. Mr. AF. Wilkinson, who was superintendent of Millpond Plantation for 40 years,
says in a letter dated October 21, 1960, that 'The original Ophiopogons, I believe, came from
France when the gardens were first landscaped some 54 years ago.' This would place the
planting at Millpond in 1906 or thereabouts" (Hume 1961, p. 150). The holotype is housed at
FLAS and a digital image! can. be seen, on the FLAS herbarium, website.

Stolons creeping, slender (fide protologue), but apparently without stolons in at least most
horticultural forms. Roots with or without tuberous swellings. Leaves (17- fide Fantz, 30-)50 -80
cm long, (5— )8— 12(— 16 fide Fantz) mm wide, thicker and less arching than in other liriopes, 7-13(-17
fide Fantz)-nerved, margins minutely serrulate distally, apices acute to obtuse to rounded. Scapes
erect, 20^2 cm long, mostly shorter than the leaves, subterete to slightly flattened, especially
distally, purplish. Inflorescence ca. 40-180-flowered, rachis 6-17 cm long, narrowly columnar with
distinctly separated fascicles; bracts lanceolate to linear, proximal 4-5 mm long. Flowers mostly in
fascicles of (2-)4-8; pedicels 3-5 mm long; tq^als light to dark violet, oblong-ovate, 4-6 mm long;
floral tube 0.3-1 mm long or less commonly essentially absent Fruits subglobose, 5-8 mm in diam.,
blue-black to dark or blackish purple at maturity. Seeds subglobose, 4-6 mm in diam. 2n =
unknown.

Native distribution unknown (see comments below and those regarding the TYPE). Common
names: GIANT LILYTURF, GIANT LIRIOPE. Representative cultivars: Aztec Grass, Emerald
Goddess, Evergreen Giant, Love Potion No. 13, Merlon Jacobs, Supergreen Giant. Fantz (2008b)
notes that "Variegated Giant," sometimes claimed to be a horticultural variant of L. gigantea, is
actually an Ophiopogon (O. jaburan). Flowering Jun-Oct. As noted by Hume (1961, p. 150):
"flowers produced earlier in the season than those of any other species growing under same
conditions; fruit also ripening ahead of that of L. muscari, L. spicata, L. graminifolia. and [L.
graminifolia]."' Broussard (2007), however, noted that it flowers June through September; cultivars
in Texas were flowering through September, essentially contemporary with L. graminifolia, mostly
later thanL. spicata.

The description above of Liriope gigantea is drawn mostly from plants seen by the author
sold and cultivated as 'giant liriope' (and usually identified as ''Liriope gigantea''''), augmented mostly

Nesom: Uriopeand Ophiopogomn the USA

for fruits from the description by Fantz (2008b). Comments by Fantz (2008b, p. 346) are useful:
'The leaves are stiff, mostly erect with some spreading apically, but not becoming arched back and
pointed downward toward the ground as observed in most species of liriopogons. The leaves are the
longest and the most elegant green of liriopogons used in the landscape/' I find that L. gigantea is
eaily distinguished from L. muscari, but the distinction is subtle and relies mostly on leaf length and
thickness.

In the protologue of Liriope gigantea (Hume 1961) it was described as a "rhizomatous plant
with stout, widely spreading rhizomes, in time forming a dense matted turf." The type specimen at
FLAS (cited above), as well as the photo in the protologue (Hume 1961, Fig. 55. p. 147) clearly show
slender, creeping rhizomes. Broussard (2007) studied samples of Liriope gigantea and also described
the root system as "rhizomatous with stout, widely spreading rhizomes." Photos of her vouchers
("Evergreen Giant" Fig. 4.7; Merton Jacobs "Supergreen" Fig. 4.19) show rhizomatous (=
stoloniferous) plants. The patent application for "Merton Jacobs" (Jacobs 2001) notes that it grows
"by adding bibs or small clumps of leaves on short rhizomes on the side of the existing clump. . . .
The rhizomes are 1/2 to 3/4 inches long. 1/4 to 1/2 inches wide. . . . [The original of the new cultivar
was] "found as an openly pollinated seedling of Liriope gigantea maintained by Flowerwood
Nursery, Inc. in Bushnell, Fla." (Jacobs 2001).

In contrast, Fantz's description noted that "rhizomatous stems lacking" is characteristic of the
growth habit of Liriope gigantea, and commercial information generally describes plants with a
clumping habit, sometimes reaching three feet in width from lateral growth. All of those I have seen
also appear to be distinctly caespitose, without rhizomes (see photos in Online Appendix). Thus it
appears that the spreading growth habit is variable in derivatives of typical L. gigantea. See
comments under L. muscari, in which similar variation perhaps occurs.

The lack of correspondence of Liriope gigantea with any known Asian species is peculiar.
Comments by Skinner (1971, p, 348-349) are pertinent: "This, the largest-leaved of all known wild or
cultivated liriopogons, is an oddity which may possibly have reached a Georgia plantation from
France early in the century. Yet it seems not to have been reported from France. ... It may well be
of garden origin." With Fantz and others, however, the present study confirms that L. gigantea is a
distinctive entity not clearly associated with any other species cultivated in the USA and apparently
not matching any recognized species of the Chinese or Japanese flora.

If Liriope gigantea indeed is of garden origin, it seems reasonable to speculate that it has
arisen from L. muscari. Plants of L. gigantea are generally larger than those of L. muscari, usually
considerably so, but large-leaved plants of L. muscari sensu lato may be similar in aspect to L.
gigantea, overlapping in leaf length and width (e.g., see the variant from south-central China in Figs.
7 and 8). Leaves of L. gigantea are thicker and more rigid (also as noted, by Fantz 2008b) and the
flowers are larger. This set of "gigas" features could be associated with an elevated ploidy level.
Variation in habit (stolons present or absent) suggests the possibility that distinct cultivars may have
arisen independently. Hybrid origin is not implausible — all plantings I have seen (Texas and
Arkansas) have been completely sterile, with no fruit production.

4. LIRIOPE SPICATA Lour., Fl. Cochinch. 1: 201. 1790 (non Convallaria spicata Thunb. 1784,
based on a type from Japan; see comments under L. exiliflora). TYPE: probably China.
"Habitat frequenter culta, incultaque in Cochinchina, & China" (holotype: P, photograph in
Bailey 1929, Figure 6).

Ophiopogon gracilis Kunth, Enum. PI. 5: 298. 1850. TYPE: Vietnam and China. As synonym fide
Tamura (in press).

Nesom: Uriopeand Ophiopogomn the USA

Ophiopogon fauriei H. Lev. & Vaniot, Repert. Spec. Nov. Regni Veg. 5: 283. 1908. Mondo fauriei

(H. Lev. & Vaniot) Farwell, Amer. Midi. Naturalist 7: 43. 1921. TYPE: Korea. As synonym

fide Chen and Tamura (2000b), Tamura (in press).

Liriope spicata vm. prolifera Y.T. Ma, J. Wuhan Bot. Res. 3: 27. 1985. TYPE: China. As synonym

fide Chen and Tamura (2000b), Tamura (in press).

Liriope spicata var. humttis F.Z. Li, Bull. Bot. Res., Harbin 6: 170. 1986. TYPE: China. As synonym

fide Chen and Tamura (2000b). Tamura (in press).

Stolons creeping, slender. Roots with tuberous swellings. Leaves (15-)25-60 cm long, 3-
6(-8) mm wide, flexuous and arching, 5-11-nerved, margins serrulate, apices acute. Scapes erect,
12-29 cm long, shorter than the leaves, terete to slightly flattened, especially distally. Inflorescence
45-72-flowered, rachis 2-5(-8) cm long, narrowly columnar with distinctly separated fascicles;
bracts lanceolate to linear, 4-6 mm long. Flowers in fascicles of (l-)2-4(-6); pedicels ca. 2 mm
long; tepals white to very pale violet, oblong-elliptic to oblong, 4.5-5 mm long; floral tube 0.5 mm
long. Fruits globose, 7-9 mm in diam., blue-black to dark or blackish purple at maturity. Seeds
subglobose, 5-6 mm in diam. 2n = 36, 72, 88, 90, 108.

Native to China, Taiwan, Japan, Korea, and Vietnam. Native habitats include forests, grassy
slopes, hillsides, moist places; ca, 10-1800 meters elevation. Common names: CREEPING LIRIOPE,
CREEPING LILYTURF, TABLE GRASS. Representative cultivars: Cassidy, Ever Blue, Franklin Mint
(or =L. muscartt), Small Dragon, Silver Dragon (= Gin Ryu) (v), Small Green. Flowering (Jul-
)Aug-Sep(-Oct).

Naturalized distribution in the USA. Alabama (Spaulding et al. 2010). Arkansas (reported
by Smith 1988 as Ophiopogon jaburan, but ID'ed here as L. spicata. Jefferson Co.: Pine Bluff,
invading vigorously in a yard, 16 Aug 1983, Locke 5235, UARK digital image:). Florida (Wunderlin
& Hansen 2008; voucher for occurrence in Wakulla Co., Anderson 24303, VDB). Louisiana (Caddo
Par.; MacRoberts et al. 2008, identified in the original report as L. muscari). Maryland (Calvert Co.:
Scientists Cliffs, Port Republic, occasional escape along Steve Easter Trail, E of picnic table, colonial
herb, 12 Oct 1986, Hill 1 7478 (GH). Mississippi. Illinois, Texas (see notes below for Texas).

Liriope spicata is a distinctive species identified with certainty by its rhizomatous habit,
narrow, arching leaves, and short scapes bearing inconspicuous inflorescences usually about mid-
level among the leaves. The flowers are white to very pale violet and relatively few pel' rachis.

i the name Liriope spicata.

In making ill.. lumiencLitut i) <.onib)nafiun Ophiopogon spicatus, Ker Gawler (1821) listed
both Liriope spicata Lour, and Convallaria spicata Thunb, as synonyms, thus the name is invalid and
the basionym for his combination is ambiguous — the listing here with Thunberg's basionym is
arbitrary. He noted that "The synonym from the 'Flora Cochinchinensis" was kindly communicated
by Mr. Brown, who has ascertained it from the original sample in the Herbarium of the Museum of
Natural History at Paris; where a portion of Loureiro's Botanical Collection had found its way from
Lisbon during the pillage of the late wars." The illustrated plant (plate 593), however, clearly is not
L. spicata Lour. It probably represents L.. muscari or L. graminifolia — although the description
notes "radice repente," the basal parts are not illi'sliated thus its idcntfh is unambiguous The
illustrated leaves are broader man characteristic of L. graminifolia but no measurements are given in
the description. Ker Gawler cited "Nepal, Cochinchina, China, and Japan" as the native range of the
species, apparently incorporating elements from the protologues of both Loureiro and Thunberg. He
noted that the illustrated plant was introduced in 1820 by the Horticultural Society of London and
grown in their conservatory — with only the generalized statement as an indication of its native
provenance.

Nesom: Liriope and Ophiopogomn the USA

Hooker (Bot. Mag. 88: t. 5348. 1862) provided an illustration and commentary associated
with the name Ophiopogon spicatus, and the name sometimes has been cited as "O. spicatus (Thunb.)
Hook. 1862." Hooker, however, clearly did not intend to provide a new name, as he cited O. spicatus
"Gawl." The illustration in Hooker's contribution probably is of L. muscari (the basal parts are not
shown or described), and regarding the previous illustration in Ker Gawler's article, Hooker noted
that "Gawler's artist has not done justice to the plant in his figure" (which also does not illustrate
basal parts).

Liriope spicata naturalized in Texas.

The occurrence of Liriope spicata has been attributed to Texas (Kartesz 2010; USDA, NRCS
2010) on the basis of a report by Neill and Wilson (2001). This Texas report, however (p. 1093),
described the species as one of several "found in varying stages of persistence at an old abandoned
home sites and cemeteries ... and not appearing to be adventive;" fee label explicitly notes that the
specimen was collected from an "Abandoned old home site ... persisting after cultivation,
rhizomatous -spreading" (Madison Co., 15 Jun 1997 [no flowers or fruits], Neill 672, TAMU digital
image!). The plants are clearly stoloniferous, with stiffly arching leaves 10-19 cm long and 2-3 mm
wide — a collection of Ophiopogon japonicus rather than L. spicata.

1. Texas. Tarrant Co.: city of Fort Worth, Overton Park between the TCU campus and Hulen Street
Overton Creek parallel to Shady Creek Drive, immediately adjacent to the creek on the sandy terrace,
10 Oct 2010, Nesom FW 10-21 (BRIT, MO, NCU, TAMU, TEX, USF).

The plants occur in at least eight large, dense, rhizomatous, well-separated colonies over a stretch of
about 200 meters (see photos in Online Appendix) along Overton Creek in Overton Park, near the
center of the city of Fort Worth. The creek at this point runs through a broad and wooded but
relatively steep-sided ravine and is about three miles above confluence with the Trinity River. The
liriope colonies are on the first terrace of the creek.

It is possible that the first plants of Liriope spicata along Overton Creek were from a local
homeowner who dumped lawn refuse down the steep banks. The creek commonly rises beyond its
banks in heavy rain and, once established in the lower terrace, a colony could be fragmented in fast

water and parts of it carried downstream.

2. Texas. Hardin Co.: Big Thicket National Preserve, Lance Rosier Unit, on the long east fork of
Rosier Park Road (Cotton Road) at the turnaround at the road end near Little Island Bayou,
bottomland hardwoods, 21 Aug 2004, Brown 30092 (SBSC, photocopy!).

Another collection from Hardin County was rq^orted by Brown et al. (2005) as Ophiopogon
jaburan — the voucher for this collection is presently unavailable for study, but it is very likely to be
Liriope spicata (in the Big Thicket National Preseve, along the Kirby Nature Trail near Turkey Creek
"far from any house or farm land," Johnson 1797, RICE transferred to TAES). Diggs et al. (2006)
included O. jaburan as a member of the East Texas flora on the basis of the report by Brown et al.

Liriope spicata is described on some internet sites as forming an "outstanding weed-
suppressing mat," an accurate description also reflective of its behavior in the Fort Worth creekside
habitat. Fruits in L. spicata are produced at a very low frequency (pers. observ.), however, which
probably limits its wider dispersal and invasive potential.

Nesom: Uriopeand Ophiopogomn the USA

Other species of Liriope.

Liriope includes a total of 6-10 species, depending on the taxonomic intepretations of L.

gigantea, L. platyphylla, and L. koreana, and L. tawadae. Brief notes on remaining taxa are provided

Liriope minor (Maxim.) Makino, Bot. Mag. Tokyo 7: 323 1893. Ophiopogon spicatus (Thunb.) Ker
Gawl. var. minor Maxim., Bull. Acad. Imp. Sci. Saint-Petersbourg 15: 85. 1870. Liriope
graminifolia (L.) Baker var. minor (Maxim.) Baker, J. Linn. Soc, Bot. 17: 500. 1879. Liriope
spicata Lour, var. minor (Maxim.) C.H. Wright, J. Linn. Soc, Bot. 36: 80 1903. Ophiopogon
minor (Maxim.) Masam., Sci. Rep. Kanazawa Univ., Biol. 5: 111. 1957. TYPE: Japan.
Kanagawa Prefecture. Yokohama, 1862, C.J. Maximowicz (probable isoiypes: GH!, without
collection number, NY digital image!; with the same locality and date but labeled
Maximowicz 1874 (P digital image!).
Liriope cernua (Koidz.) Masam., Mem, Fac. Sci. Taihoku Imp, Univ. 11: 559. 1934, Mondo cernuum
Koidz., Bot. Mag. Tokyo 40: 332. 1926. TYPE: Japan. As synonym oft, minor fide Tamura
(in press).
Mondo tokyoense Nakai, J. Jap. Bot. 19: 314. 1943. TYPE: Japan. As synonym ofL. minor fide

Tamura (in press).
Liriope angustissima Ohwi, Acta Phytotax. Geobot. 3: 201. 1934. Liriope minor (Maxim.) Makino
var. angustissima (Ohwi) S.S.Ying, Fl. Taiwan ed. 2, 5: 53. 2000. TYPE: Taiwan. As
conspecific with L. minor fide Ying (2000); as synonym of L. graminifolia fide Chen and
Tamura (2008a).
Liriope minor was included in the overview by Fantz (2008b, p. 347), who noted that it is "a rare
species in the green industry, found in limited botanic gardens labeled as a cultivar of Liriope or
Ophipogon japonicus." It is stoloniferous-colonial and further unequivocally characterized by its
short and narrow leaves and especially by its scapes shorter than the leaves and with short, very few-
flowered rachises. The native range of L. minor is China, Japan, Korea, and the Philippines. Chinese
plants of the species are described with leaves 2-3(-4) mm wide and 7-20 cm long, scapes 6-7(-15)
cm long with rachises l-3(-5) cm long and 5-12 flowers (Chen & Tamura 2000a); in Japan, the
leaves are 8^12 cm long, 2-3 mm wide, scapes 3.5-12(-16) cm long with rachises 0.5-5(-9) cm long
and 3-17 flowers (Tamura in press). Analogous measurements for L. minor by Fantz indicate that his
concept of the species is similar: leaves 2-4 mm wide, (5.5-)7.5-15 cm long, scapes 5-8.5 cm long,

jikI l.ljll^^ I _' ^ .111 I.'IIV U llll ,..i ' IN 11. -uc^

Liriope koresma (Palib.) Nakai, Bot. Mag. Tokyo 48: 777. 1934. Ophiopogon spicatus var. koreanus

Palib., Trudy Imp. S.-Peterburgsk. Bot. Sada 19: 106. 1901. Liriope graminifolia (L.) Baker

var. koreana (Palib.) Nakai, Fl. Kor. 2: 239. 1911. Mondo koreanum (Palib.) Hatus., Bull.

Kyushu Imp. Univ. Forests no. 5: 231, 1934. Liriope spicata Lour, forma koreana (Palib.) H.

Hara, J. Jap. Bot. 59: 38. 1984. TYPE: Korea.

Liriope koreana has been treated as a synonym of L. spicata by Chen and Tamura (2000b) and

Tamura (in press), but in the limited material I have seen (GH: Japan, Shinsyu; Korea, Sorai Beach,

Whanghai), plants identified as L. koreana do not appear to be the same species as L. spicata. The

scapes are equal to or longer than the leaves and produce more and smaller flowers than in L. spicata,

and the leaves are 10-20 cm long and 2-4 mm wide.

i (Batalin) C.H. Wright, J. Linn. Soc, Bot. 36: 79, 1903, Ophiopogon ka
Batalin, Trudy Imp. S.-Petersburgsk Bot. Sada 13: 103. 1893. Mondo kansuense (Batalin)
Farwell, Amer. Mdl. Naturalist 7: 42. 1921. TYPE: China.
Liriope kansuensis is endemic to a relatively small region of central China, in northern Sichauan
Province and adjacent Gansu Province. It is distinctive in its stoloniferous habit, short linear leaves,
and few-flowered scapes longer than the leaves.

Nesom: Uriopeand Ophiopogomn the USA

Liriope pedicellata Wang & Tang, Fl. Reipubl. Popularis Sin. 15: 251. 1978. TYPE: China.

Liriope pedicellata occurs only at moderate elevations in northeastern Sichuan Province. The plants
are characterized by their caespitose habit, long, linear leaves, and as the epithet implies, the
distinctively long pedicels.

Liriope tawadae Ohwi, Acta Phytotax. Geobot. 5: 55. 1936. Ophiopogon tawadae (Ohwi) Masam.,

Sci. Rep. Kanazawa Univ. 5(a): 111. 1957. TYPE: Ryukyu Islands (Japan, Okinawa).
Tamura (in press) characterizes Liriope tawadae as an endemic of the Ryukyu Islands and
differentiates it from L. muscari by its angled scape and longer pedicels articulated near the apex —
but entering the caveat "Further studies are needed in order to clarify whether Liriope tawadae is
really distinguished from L. muscari or not."

5. OPHIOPOGON JAPONICUS (Thunb.) Ker Gawl., Bot. Mag. 27, plate 1063. 1807. Convallaria
japonica Thunb., Nova Acta Regiae Soc. Sci. Upsal. 3 [Kaempferus Illustratus I]: 208. 1780
[also Fl. Jap., 140. 1784]. Flueggea japonica (Thunb.) Rich., Neues J. Bot. (Schrader) 2: 9.
1807. Slateria japonica (Thunb.) Desv., Verh. Batav. Genootsch. Kunsten 12: 15. 1830.
Mondo japonicum (Thunb.) Farwell, Amer. Midi. Naturalist 7: 42. 1921. TYPE: Japan, [C.P.
Thunberg 8491] (holotype: UPS, photo in Bailey 1929 as Fig. 5).

Authorship of Convallaria japonica is sometimes attributed to L. f (Suppl. PI., 204.
1782), but Thunberg first published the ume Kti Gawlei u f u! lhuisb^ig s publication in
Flora Japonica, but the description in Kaempferus Illustratus was two years earlier. Linnaeus
cited '"Habitat in Japonia. Thunberg." Ker Gawler augmented his description of the species
with information from Kaempfer's Amoenitatum Exoticarum of 1712.

Convallaria japonica var. minor Thunb., Fl. Jap. 139. 1784. TYPE: Japan, As synonym fide Chen
and Tamura (2000b), Tamura (in press).

Ophiopogon stolonifer H. Lev. & Vaniot, Mem. Accad. Pontif. Nuovi Lincei 23: 344. 1905 [Liliac.
etc Chine, 16. 1905]. Mondo stolonifer (H. Lev. & Vaniot) Farwell, Amer. Midi. Naturalist
7:42. 1921. TYPE: China. As synonym fide Chen and Tamura (2000b), Tamura (in press).

Stolons creeping, slender. Roots with tuberous swellings. Leaves (5-)12-35(-40) cm long,
2-4 mm wide, 3-5(-7)-nerved, margins serrulate, apices acute to obtuse or rounded. Scapes erect, 6-
15(-27) cm, much shorter than iea\e> UightK flattened Inflorescence ca. 5-10+-flowered, rachis
2-5 cm; bracts lanceolate, proximal 7-8 mm long. Flowers usually nodding, in fascicles of l-2(-3);
pedicels 2-2.5 mm, articulate near middle; tepals white to light lilac or pale purple, lanceolate, 3.5-
4.5 mm long; floral tube 1-2 mm long. Fruits subglobose to ellipsoid-globose, ca. 7-9 mm in diam.,
blue to blue-violet at maturity. Seeds globose, 6-8 mm in diam., blue at maturity (photos in the
Online Appendix). 2n = 34, 36, 67, 68, 72, 108.

Native to China (incl. Taiwan). Japan, and Korea. Native habitats include forests, dense
scrub in ravines, and moist and shady places on slopes and along streams, cliffs; 200-2800 meters
elevation. Widely cultivated in China for its tuberous roots, which are used medicinally. Common
names: MONDO GRASS, MINI MONDO, DWARF MONDO, SUPER DWARF MONDO, ENGLISH
MONKEYGRASS. Representative cultivars: Albus, Bluebird, Caeruleus, Comet (v), Compactus,
Gyoko Ryu, Jade Dragon, Kijimafukiduma (v), Kyoto Dwarf, Minor, Nanus, Nippon, Silver Mist (v,
= Kijimafukiduma?), Silver Showers (v), Shiroshima Ryu, Striatus, Super Dwarf, Tears of Gold.
Torafu. Several of the cultivar names refer to plants with extremely abbreviated leaves (e,g.,
Compactus, Gyoko Ryu = Super Dwarf, Kyoto Dwarf, Nanus = Dwarf), Flowering May-Jun;
flowers apparently produced uncommonly.

Naturalized distribution in the USA: Alabama (fide Kartesz 2010). Diggs et al. (1999) noted

that Ophiopogon japonicus in north central Texas is widely cultivated and persists and spreads

Nesom: Uriopeand Ophiopogomn the USA

vegetatively. A photo on the Bayou Preservation Association website <http://www.bayou
preservation.org/> shows fruits of O. japonicus and the species is included among "The Invasive Exotic
"Dirty Dozens" — presumably the photograph was taken of a naturalized plant somewhere in the
bayou area around Houston in southeastern Texas but no documentation is provided. See Spaulding
et al. (2010) for another photo of a fruiting plant of O. japonicus in a naturalized population. I have
rarely seen fruits in cultivated plants.

Differences can be striking between longer-leaved horticultural forms and the dwarf, highly
compact forms with leaves as short as 5 centimeters. The smaller plants md dwarfs are derived from
the longer-leaved wild type.

Tamura (in press) recognizes three varieties within Ophiopogon japonicus. Var. caespitosus
Okuyama (endemic to Japan) and var. umbrosus Maxim. (Japan. Korea, and China) both are
characterized by stolons much reduced in length (compared to var. japonicus) and more vigorously
branching rhizomes. Presumably the forms seen in USA horticulture all are var. japonicus, since all
appear to produce creeping stolons and are strongly colonial.

6. OPHIOPOGON PLANISCAPUS Nakai, Cat. Sem. Hort. Tokyo 20. 1919; 33. 1920. Mondo
planiscapum (Nakai) L.H.Bailey, Gentes Herb. 2: 25. 1929. TYPE: Japan.

Stolons usually creeping, slender, sometimes shorter. Roots with tuberous swellings. Leaves
15-30(-50) cm long (fide Tamura in press; apparently mostly less than 25 cm in popular cultivars),
4-6(-7) mm wide, 7-11-nerved, margins serrulate distally apices obtuse. Scapes erect or arching, ca.
10-20 cm long, shorter than the leaves, flattened, winged, sometimes weakly 3-angled near the base.
Inflorescence 10-32-flowered, raehis 4-8 cm long, often 1-sided; fascicles (5-)12-18; bracts
lanceolate, proximal 8-18 mm long. Flowers pendulous, in fascicles of 1-3; pedicels 2-6 mm long;
tepals pale purple to pinkish or white, 4.5-5.5 mm long; floral tube 2-4 mm long. Fruits ellipsoid-
globose, 5-6 mm in diam., dark blue to blue-black or purple-gray at maturity. Seeds globose, 8-9
mm in diam., dark blue to purplish black or black at maturity. 2n = 36.

Native and endemic to Japan. Native habitats are forest floors and forest margins; ca. 50-
1000 meters elevation. Common names: BLACK MONDO GRASS, BLACK MONDO, BLACK
LILYTURF. Representative cultivars: Arabicus, Black Dragon, Dark Beauty, Ebony Knight,
Leucanthus, Little Tabby (v), Minimus, Nigrescens, Silver Ribbon (v), Silver Shine (v). Flowering
(Jun-)Jul-Aug.

Not reported in the USA outside of cultivation.

Ophiopogon planiscapus is recognized by its large, white to lilac-tinged, campanulate flowers
in fascicles of 1-3 and especially by its relatively narrow leaves that become (in the popular
horticultural forms) dark purple to black (Howe 2004). Wild type leaves apparently are dark green
(see photos in Online Appendix).

7. OPHIOPOGON JABURAN (Sieb.) Loddiges, Bot. Cab. 19, plate 1876. 1832. Slater ia jabur an

Sieb., Verh. Batav. Genootsch. Kunsten 12: 15. 1830. Flueggea jaburan (Sieb.) Kunth,
Enum. PI. 5: 303. 1850. Mondo jaburan (Sieb.) L.H. Bailey/ Gentes Herb. 2: 27. 1929.
TYPE: Japan. Nagasaki: Nomo-Saki, 1863, C.J. Maximowicz s.n. (holotype: presumably LE;
probable isotype: P digital image!, with cited collection information).

A photo of a UPS Thunberg specimen in Bailey (1929, Fig. 3; Japan, "Convallaria
japonica a") was noted by him as representative of Ophiopogon jaburan.

Nesom: Uriopeand Ophiopogomn the USA

Convallaria japonica L. f. var. major Thunb., Fl. Jap., 139. 1784. Flueggea japonica (L. f.) Rich.

var. major (Thunb.) Schult. & Schult. f. in Roemer & Schultes] Syst. Veg. 7: 309. 1829.

TYPE: Japan. As synonym fide Tamura (in press).
Ophiopogon taquetii H. Lev., Repert. Spec. Nov. Regni Veg. 8: 171. 1910. Mondo taquetii (H. Lev.)

Farwell, Amer. Midi. Naturalist 7: 43. 1921. TYPE: Korea. As synonym fide Tamura (in

press).

Stolons absent or very short and vertically oriented. Roots with tuberous swellings. Leaves
linear, 30-85(-130) cm long, (7-)10-18 mm wide, 9-13(-17)-nerved, upper margins minutely
serrulate, apices acute. Scapes arching, 24-40(-75) cm long, shorter to longer than the leaves,
strongly flattened, winged at margins. Inflorescence 45-120-flowered, rachis 6— 10(— 15) cm long,
often 1-sided; fascicles 12-20; bracts narrowly lanceolate to lanceolate, proximal 15— 85(— 180) mm
long. Flowers in fascicles of 3-8C-10); pedicels 4-6(-8) mm long; tepals usually white, rarely
purplish, ovate-oblong, 5-7 mm long, slightly recurved; floral tube 3-6(-8) mm long. Fruits oblong,
10-15 mm long, violet-blue to dark violet at maturity. Seeds ellipsoid to oblong, (5-)8-14 mm long,
white to deep blue at maturity. In = 36.

Native to Japan, Korea, and Taiwan. Native habitats are primarily forests in coastal
localities; ca. 5-50 meters elevation. Common names: GIANT LILYTURF, WHITE LILYTURF, AZTEC
GRASS, SNAKEBEARD, JAPANESE HYACINTH, JABURAN LILY. Representative cultivars: Argenteo-
variegatus (v), Aureo-variegatus (v), Caeruleus, Crow's White. Javanensis, Variegatus (v), Vittatus
(v), White Dragon (v). Flowering Jun-Aug.

Probably not naturalized in the USA; see comments under Liriope spicata.

According to Fantz (2009, p. 390), "variegated selections [of Ophiopogon jaburan] are most
commonly used in interior landscapes and conservatories. Outside [in North Carolina], the variegated
selections usually did not survive into the second season."

8. OPHIOPOGON INTERMEDIUS D. Don, Prodr. Fl. Nepal. 48. 1825. Flueggea japonica var.

intermedia (D. Don) Schultes. Syst. Veg. 7: 310. 1829. Ophiopogon japonicus var.

intermedins (D. Don) Maxim., Melanges Biol. Bull. Phys.-Math. Acad. Imp. Sci. Saint-

Petersbourg 7: 327. 1870. Mondo faponicum var. intermedium (D. Don) Farwell, Amer.

Midi. Naturalist 7: 42. 1921. Mondo intermedium (D. Don) L.H. Bailey, Gentes Herb. 2: 25.

1929. TYPE: Nepal.
Flueggea wallichiana Kunth, Enum. PI. 5: 303. 1850. Ophiopogon wallichianus (Kunth) J.D. Hook.,

Fl. Brit. India 6(18): 268. 1892. Mondo japonicum var. wallichianum (Kunth) Farwell, Amer.

Midi. Naturalist 7: 42. 1921. Mondo wallichianum (Kunth) L.H. Bailey, Gentes Herb. 2: 21.

1929. TYPE: India (Silbet). Possible type: without collection data, N. Wallich 5139 (PH

digital image!). As synonym fide Chen and Tamura (2000b).
Flueggea jacquemontiana Kunth, Enum. PI. 5: 304. 1850. TYPE: "India orientalis," As synonym

fide Chen and Tamura (2000b).
Flueggea dubia Kunth, Enum. PI. 5: 305. 1850. TYPE: "India orientalis." As synonym fide Chen

and Tamura (2000b).
Flueggea griffithii Baker, J. Linn. Soc, Bot. 17: 502. 1879. Ophiopogon griffithii (Baker) J.D.

Hook, Fl. Brit, India 6(18): 270. 1892. Mondo japonicum var. griffithii (Baker) Farwell,

Amer. Midi. Naturalist 7: 42. 1921. TYPE: India ("Himalaya orientalis in sylvis ad

Patkaye"). As synonym fide Chen and Tamura (2000b).

Stolons creeping, long or short. Roots with tuberous swellings. Leaves 15-55(-70) cm
long, 3-15 mm wide, 5-9-nerved [13-23-nerved fide Fantz 2009], margins usually serrulate, apices

Nesom: Uriopeand Ophiopogomn the USA

Scapes projecting outward, (12-)20-50 cm long, shorter than or equalling the leaves, 3-angled
flattened and narrowly 2-winged. Inflorescence ca. (100-)200-300-flowered, rachis
(2.5-)5-16 cm long [(8-)17-21 cm long fide Fantz]; fascicles (30-)45-55; bracts subulate, linear, or
lanceolate, proximal 15-40 mm long. Flowers spreading, in fascicles of (l-)2-3(-6); pedicels 2-3
mm; tepals white to light purplish, oblong, ovate, or ovate-lanceolate, 4-7(-9) mm long; floral tube
2.5-3 mm long. Fruits oblong to ovoid to ellipsoid-globose, 8-11 mm long, 7-8 mm in diam,
blackish purple to blue at maturity. Seeds ellipsoid, ca. 4 mm in diam., blue. 2n = 36, 54, 68, 72,
108, 112.

As measured from plants cultivated at the University of Texas (see photos in Online
Appendix). Leaves 23-40 cm long, 4-7 mm wide, 10-13-nerved, upper margins minutely serrulate,
apices acute. Scapes arching-spreading, 8-14 cm long, shorter than the leaves, strongly flattened.
Inflorescence 50-75 flowered, rachis 8-10 cm long; bracts ovate-lanceolate, proximal 4-5 mm long.
Flowers in fascicles of 2-4(-6); pedicels 2.5^1 mm long; tepals white, oblong to oblong-lanceolate,
4-6 mm long; floral tube 1.5-2 mm long.

Native to the Himalayan region (India, Nepal, Bhutan, southwestern China) and Sri Lanka, to
Bangladesh, Myanmar, Thailand, Vietnam, and Taiwan, Natr\ ~ h i^itats include e\ ei g een broad-
leaved forests, mixed forests, pine forests, bamboo forests, scrub, moist and shady places along
streams, grassy slopes; 700-3000 meters elevation. Common names: HIMALAYAN LILY LURF,
WHILE MONDO GRASS, AZLEC GRASS. Representative cultivars: Argenteomarginatus (v),
Compactus, Stripey White. Flowering Jun-Oct.

Not reported in the USA outside of cultivation.

Ophiopogon intermedins lias one of the most widespread native geographic ranges in the
genus and shows high levels of genetic diversity (allozymes) as well as morphological and
chromosomal variation (He et al. 2000). Within the genus, O. intermedins and O. planiscapus
apparently are closely related (Tanaka 2001-III).

According to Fantz (2009, p. 390), Ophiopogon intermedins is "marketed commonly under
Liriope mnscari 'Variegated Evergreen Giant' or 'Grandiflora White.' Also, this species is confused
with variegated selections of O. jaburan." The species is described by Chen and Tamura (2000b) as
producing short, thick rhizomes; Fantz (2009) observed that it produces rhizomes (stolons) 7-28 cm
long and forms dense tufted mounds with age. Fantz also noted that dwarf nursery selections of O.
intermedins are 12-18 cm tall.

ACKNOWLEDGEMENTS

I am grateful for the arrangement of a loan of Liriope from GH to TEX, to David Boufford
(Harvard Herbaria) for information on Japanese species, Amber Schoneman (at TEX) for providing
digital images of the GH specimens included in the manuscript as Figs. 3-8, Monique Reed (TAMU)
for sending an image and observations on the collection from Madison Co., Texas (see comments
under L. spicata), Larry Brown (SBSC) for sending a photocopy of his liriopogon collection (see L.
spicata) from the Big Thicket National Preserve, Michael and Barbara MacRoberts (LSUS) for
information and images of Louisiana plants, Johnnie Gentry and Brent Baker for sending an image of
the collection of L. spicata naturalized in Arkansas, to Lowell Urbatsch (LSU Botany), John Praski
(MO), Marie Long (NYBG Library), and Mary Stiffler (MBG Library) for sending pertinent
literature, to Nancy Elder (University of Texas Life Sciences Library) for on-site help with literature,
and to Paul Fantz for a liriopogon discussion and for sending his publications on these plants. Many
thanks to Dr. Mats Hjertson, UPS Curator, for sending digital images of the type specimen of
Convallaria spicata Thunb.; Peter Nielson, owner of Ace Mondo Ltd., for images of Ophiopogon

Nesom: Uriopeand Ophiopogomn the USA

planiscapus and permission to use other photos posted on his website; Glenn Kopp, Horticultural
Information Manager at the Missouri Botanical Garden, for permission to use a photo; Catherine
Broussard to permission to use photos and for comments on the manuscript; Michael Charters of
Calflora for permission to use photos; and Robert George (BRIT), who discovered the naturalized
plant of L. graminifolia in Euless, Texas, and took me to see it.

LITERATURE CITED

Adams, G. 1989. Great ground covers. Amer. Nurseryman 170: 83-91.

Ahmad, V.U. and A Basha (eds.). 2006. Spectroscopic Data of Steroid Glycosides. Vol. 2.

Springer, New York.
Bailey, L.H. 1924. Manual of Cultivated Plants. Macmillan Co., New York.
Bailey, L.H. 1929. The case of Ophiopogon and Liriope. Gentes Herb. 2: 1-37.
Bailey, L.H. 1949. Manual of Cultivated Plants (rev. ed.). Macmillan Co., New York.
Bailey Hortorium Staff. 1976. Hortus Third: A Concise Dictionary of Plants Cultivated in the United

States and Canada. Liberty Hyde Bailey Hortorium, Cornell Univ., Ithaca, New York.
Baker, J.G. 1875. Revision of the genera and species of Asparagaceae. J. Linn. Soc, Bot. 14: 508-

632.
Broussard, M.C. 2007. A horticultural study of Liriope and Ophiopogon: Nomenclature,

morphology, and culture. Ph.D. dissertation, Louisiana State Univ., Baton Rouge.
Brown, L.E., B.R. MacRoberts, M.H. MacRoberts, P. Harcombe, W.W. Pruess, S. Elsik, and D.

Johnson. 2005 . Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big

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i: Uricpeand Ophiopogon in the USA 24

Figure 1 . Holotype of Convallaria spicata Thunb. at UPS. identified here as Liriope r,
(Dcne.)L.H. Bailey.

Nesom: Liriope and Ophiopogon'm

r. Liriope and Ophiopogon'm th

Figure 3. Liriope muscari, typical expression. Note the short, thickened rhizome, apparently n
commonly characteristic of this species in Japan than elsewhere.

Figure 4. Liriope muscari, typical expression. Base apparently strictly caespitos

im: Ur/opeand Ophiopogon'm the USA 28

. Liriope muscari sensu lato, "platyphylla" expression (see comments in text).

u lato, "platyphylla" expression (se

id Ophiopogon in the USA 3 ]

Liriope muscari sensu lato, variant of "platyphylla" expression (see comments in text).