Content uploaded by Mark Garland
Author content
All content in this area was uploaded by Mark Garland on Jan 14, 2022
Content may be subject to copyright.
INTRODUCTION
Hymenocallis Salisb., a New World genus recently
placed together with Ismene Salisb. ex Herb. and
Leptochiton Sealy in the tribe Hymenocallideae of the
Amaryllidaceae (Meerow & al., 2000 a, b), may contain
the most difficult group of flowering plants in which to
recognize species solely by study of herbarium speci-
mens. Habit characters critical for delimitation and iden-
tification of species, such as texture, surface features and
the position of the leaves, are rarely apparent in pressed
specimens. Perhaps even more obscure are floral charac-
teristics like the shape of the staminal cup, the pattern of
dentation of the margin of this cup at full anthesis, and
the position and colour of the perianth rays. Partly
because of this difficulty, twentieth-century treatments of
the genus (Small, 1933; Morton, 1935; Sealy, 1954;
Traub, 1962) seem to have failed to describe accurately
the variation of Hymenocallis species in nature, and
workers who have attempted to identify specimens using
these treatments, or local floras based on them, general-
ly found the results unsatisfactory. In their wetland flora
of the southeastern United States, Godfrey & Wooten
(1979) did not even attempt to describe the species of
Hymenocallis in the region, citing as reasons for this the
scarcity and poor quality of herbarium specimens, the
lack of studies on populations in the wild, and the short-
comings of publications on the genus.
Beyond the difficulty of recognizing species in
Hymenocallis is the difficulty of applying correct names
to the species that are recognized. These showy plants
are attractive to gardeners, so that many of the species
now included in the genus were described from living
plants cultivated in England or other European countries,
sometimes without detailed knowledge of their place of
origin. More importantly, as Sealy (1954) pointed out,
almost no specimens of the species described between
1753 and 1853 were preserved in any herbaria. In some
cases, these species were illustrated, but in other cases,
descriptions of sometimes dubious quality are the only
evidence available to fix the application of the names.
Sealy (1954) traced the history of the species now
included in Hymenocallis both before and after Salisbury
(1812) separated the American spider-lilies from
Pancratium and established the genus Hymenocallis on
the basis of the “beautiful membrane” that forms the sta-
minal cup. Herbert (1837), Roemer (1847), Kunth (1850)
and Baker (1888) each treated all species then included
in Hymenocallis.
Small (1933) provided the first comprehensive
review of the southeastern United States species in the
20th century. We consider his treatment a valiant attempt
to define the species of the South, and Smith & Flory
(2002) recognized several of his species in their treat-
ment for the Flora of North America. Small’s narrow
species concepts and the age of his treatment, which of
course excludes later-described taxa, prevented his work
from being more useful.
Morton (1935) provided a nomenclatural treatment
of Hymenocallis. He listed thirteen species and syn-
onyms under each. Although our treatment recognizes
only five of his species names as having priority, we
found his treatment valuable in helping us to sort out the
805
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
N O M E N C L A T U R E
Edited by John McNeill & Gerry Moore
Nomenclature of Hymenocallis taxa (Amaryllidaceae) in southeastern United
States
Ger ald L . Smith1& Mark A. Garland2
1Biolog y De par tm ent, High P oint Unive rsity, Un ive rs ity Stat ion -M ontl ieu Avenue , H ig h Po int, No rth Ca ro lina
27 262, U.S.A. g smi th@ high point.ed u (au th or f or corresp on den ce )
2Botany Se ction, D iv is io n of P la nt Ind ust ry, Florid a D epartme nt o f Ag ricu lture a nd Co n su mer S er vice s, Post
Offi ce B ox 14710 0, G a in es ville, Fl orida 3 2614, U .S .A . garla nm 1@do acs .stat e.fl.us
For the Flora of North Amer ic a, Sm ith & F lo ry (20 02) reco gniz ed 1 5 sp ecies an d four inf ra spec if ic taxa o f
Hym enocallis in th e so uthe aste rn U nited Sta tes from Tex as and O k la ho m a ea st to Virgi nia. Her e w e ju stify the
sele ction of nam e s us ed for thes e taxa in the Flora o f North A meric a treatment and typify those names.
KEYWORD S: Am a ryll idac ea e, H y me n oc allis, nom enc latu re, t ypif icat ion.
names that had been described for Hymenocallis taxa up
to 1935.
Sealy (1954) presented the first worldwide treatment
of Hymenocallis since Baker (1888). This was based on
specimens at BM, E, and K, and living plants that flow-
ered at Kew during 1953. He recognized 27 species and
presented a key for their identification. For each species,
he listed synonyms, gave a synoptic description and list-
ed herbarium specimens that he had examined. He also
included information about the distribution of each
species, emphasized distinctive features and provided an
informative taxonomic discussion. Sealy nevertheless
realized that his efforts had their shortcomings and rec-
ommended the study of many more specimens, especial-
ly study of plants in the wild, in order to delimit species
more accurately.
Traub (1962) presented a synoptic key to Hymeno-
callis throughout its range and grouped the species into
alliances based on characteristics of the leaves, number
of ovules, chromosome numbers and geographical distri-
bution. For much of his knowledge of Hymenocallis taxa
of the southeastern United States he was indebted to Mrs.
Mary G. Henry of Gladwyne, Pennsylvania, who collect-
ed living plants from the Atlantic seaboard and the Gulf
Coast. Mrs. Henry shared most of her living collections
with Traub, who cultivated them and described many as
new species. Mrs. Henry’s careful documentation of
localities and habitats in her field diaries has enabled us
to revisit many of the original localities of the collections
on which Traub based his species.
However, the vagueness of Traub’s treatment dimin-
ishes its utility. He did not include all synonyms and, as
a result, taxonomists are left to wonder about the status
of such a familiar name as H. crassifolia Herb. He also
failed to provide full descriptions of the species and said
little about geographical distribution. The treatment is
also sorely lacking in ecological information such as
habitats and plant associates. According to Robert K.
Godfrey (pers. comm.), trying to use Traub’s key to iden-
tify Hymenocallis species is a rather impossible task.
Flory (1976, 1978) reported cytological information
for many taxa of Hymenocallis. The chromosome num-
bers and karyotypes recorded in these papers have
proved of great value in corroborating the morphological
distinctions used for recognizing the taxa in the treatment
of Hymenocallis for the Flora of North America (Smith
& Flory, 2002) and in this paper.
In this paper we follow Smith & Flory (2002) and
accept fifteen species of Hymenocallis as growing natu-
rally in the southeastern coastal plain and piedmont of
the United States from eastern Texas to Virginia. These
species are morphologically and cytologically recogniz-
able entities that we have distinguished after extensive
study of the plants in the field during the last 18 years,
especially in Florida where twelve of the fifteen species
occur, from examination of numerous herbarium speci-
mens and from extensive cytological analyses.
Recognition of these species does not resolve all the
problems in the taxonomy of southeastern Hymenocallis,
but it provides a basis for future studies in the genus.
MATERIALS AND METHODS
We list below the names of all species and infraspe-
cific taxa of Hymenocallis that were treated by Smith &
Flory (2002). For each taxon name we list a type. For
those names without a type designated by their author,
we have either selected lectotypes from the original
material or designated neotypes that, as closely as possi-
ble, agree with the original description of the taxon. In
some cases we have chosen recent specimens as epitypes
to represent lectotypes that are illustrations. In a number
of cases we have selected as lectotypes illustrations that
accompanied the protologue; we recognize that, under
the definition of “original material” in the ICBN (cf. Art.
9 Note 2(a) in the St. Louis Code; Greuter & al., 2000),
such illustrations will generally not constitute original
material (cf. Ross, 2002). As, in each case, this is the
only potentially original material available, we have
maintained the widespread practice of referring to these
as lectotypes, even if a strict interpretation of the Code
would in some, if not all, of these cases require the selec-
tions to be considered neotypes.
Laferriere (1996) argued that as Traub designated
multiple sheets as type of several of his species names,
they are invalid under Article 37.3 of the 1994 version of
the International Code of Botanical Nomenclature
(“Tokyo Code”; Greuter & al., 1994) which required
“citation of a single element…as indication of the holo-
type”. Laferriere therefore republished several of these
names (including Hymenocallis choctawensis, H. pun-
tagordensis,H. henryae, H. moldenkiana,H. palusviren-
sis and H. duvalensis), designating a single sheet from
amongst those cited by Traub as type for each name. We
have previously argued (Smith & Garland, 1996) that the
sheets of each of Traub’s type collections represent parts
of a single plant, and that if the sheets of each type col-
lection, taken together, were interpreted as a “single ele-
ment”, then Laferriere’s republication was not necessary.
Article 8.2 of the current edition of the ICBN, the “St.
Louis Code” (Greuter & al., 2000), states that “for the
purpose of typification a specimen is a gathering, or part
of a gathering, of a single species or infraspecific taxon
made at one time.... It may consist of a single plant, parts
of one or several plants, or of multiple small plants”, and
Article 37.2 explicitly allows a type to be designated “by
reference to an entire gathering, or part thereof, even if it
Smith & Garland Nomenclature of Hymenocallis 52 November 2003: 805–817
806
consists of two or more specimens”. We consider that
each of Traub’s types represents a single gathering, and
so retain Traub as the author of these names and use his
original type designations. The editors of the journal
Herbertia (Vassar & Meerow, 1997) agreed that
Laferriere’s republication of these names was inappro-
priate and issued a statement retracting his paper.
Under each taxon below we list all the synonyms
known to us that can be definitely assigned to that taxon.
We also briefly describe the major morphological char-
acteristics and geographic range of each taxon.
Measurements of length and width follow the format
length × width; for example, “3.5–6 dm × 2–6 cm”.
Extreme values, greater than two standard deviations
from the mean, are enclosed in parentheses: “2.5–4
(–4.5) cm”.
TAXONOMIC TREATMENT
Hymenocallis Salisb., Trans. Hort. Soc. London 1: 338.
1812. Lectotype (see Traub, Gen. Amaryllidaceae
74. 1963): Hymenocallis speciosa (L.f. ex Salisb.)
Salisb., Trans. Hort. Soc. London 1: 340. 1812 º
Pancratium speciosum L.f. ex Salisb., Trans. Linn.
Soc. 2: 73. t. 12. 1794 (superseding, under Art.
10.5(b), the choice of H. littoralis by N. L. Britton &
A. Brown, Ill. Fl. N. U.S. ed. 2. 1: 533. 7 Jun 1913).
Original locality: “Floruit hactenus in nostro cal-
dario tantum mense Octobris” [greenhouse at Chapel
Allerton, Yorkshire, England].
Description: Glabrous, perennial herbs with large,
onion-like tunicated bulb; leaves deciduous or persistent,
strap-like, often coriaceous, bright green to blue-green,
channelled, distichous, arching to erect; flowers 1-many
per inflorescence, fragrant, consisting of a white, mem-
branous, saucer-like staminal cup created by the fusion
and expansion of proximal portions of filaments with the
distal portions of filaments free and “spidery”, subtend-
ed by 6 white or pale green, narrow perianth rays extend-
ing outward from the top of a green, slender floral tube;
ovary sessile or rarely pedicellate atop a scape; fruit a
fleshy capsule with 1–6 seeds (Fig. 1).
Recognition. Bulbous, perennial herbs with fleshy,
coriaceous, strap-like, shiny green to blue-green, distic-
hous leaves; flowers with a membranous staminal cup
above 6 long and narrow, white to pale-green perianth
rays. The genus includes 30 to 40 species (Mabberley,
1997). We recognize fifteen species in the southeastern
United States. They are extremely difficult to identify as
herbarium specimens but are more easily identifiable in
the field; for an identification key to the taxa of the
southeastern United States, see Smith & Flory (2002).
The similar Crinum americanum (swamp-lily) may be
easily distinguished from spider-lilies by its flowers that
lack a staminal cup and leaves that are spirally, instead of
distichously, arranged, and have fine teeth along the mar-
gin.
1. Hymenocallis occidentalis (Leconte) Kunth, Enum.
Pl. 5: 856. 1850 ºPancratium occidentale Leconte,
Ann. Lyceum Nat. Hist. New York 3: 146. 1836 º
Tomodon pratense Raf., Fl. Tellur. 4: 22. 1838, nom.
superfl. Original locality: U.S.A.: “in the western
parts of the state of Georgia and in Tennessee and
Kentucky”; no type specimen located. – Neotype
(designated here): U.S.A. Georgia, Bartow County:
on terrace, a few feet above level of Allatoona Creek,
24 Aug 1948, W. H. Duncan 8828 (GA!)
Description: Bulb nonrhizomatous, globose. Leaves
light green, glaucous, noncoriaceous, arching to erect,
shallowly channelled, oblanceolate, distinctly wider
above the middle but tapering to a petiole-like portion
proximally, 3.5–6 dm × 2–6 cm. Flowers 3–9 per scape,
strongly fragrant; perianth rays white, 7–12 cm long; sta-
minal cup funnelform, 2.5–4(–4.5) × 4–5.5 cm, with
margin often irregularly tri- or bi-dentate between the
free portions of the filaments.
Diagnosis: Hymenocallis occidentalis possesses
oblanceolate, subpetiolate leaves, as does H. choctawen-
sis, but differs from that species by possessing glaucous
leaves and scape bracts that are lanceolate and with long-
acuminate apex.
1a. Hymenocallis occidentalis (Leconte) Kunth var.
occidentalis
=Hymenocallis bidentata Small, Man. S.E. Fl. 323,
1503. 1933. – Type: U.S.A. Alabama, Cullman
County: St. Bernard, 20 Oct 1920, Bede Knapke s.n.
(holotype: NY, fide Small [1933], p. 1503; type
specimen not located.)
=Hymenocallis moldenkiana Traub, Pl. Life 18: 71.
1962. – Type: U.S.A. Georgia, Appling County: 12
miles NE of Baxley, from bulbs collected by Mrs.
Mary G. Henry (T-223), specimen H. P. Traub 272a
and b(holotype: MO! [ex TRA]).
Description: Leaves appearing in late winter, fresh
at anthesis, then dying, arching outward, up to 6 cm
wide.
This is the most widely distributed of any
Hymenocallis species in the United States, occurring
from southwestern North Carolina to the hammocks of
the Florida panhandle westward to Louisiana and north-
ward to southern Missouri and Illinois. It inhabits flood-
plain forests, hammocks, meadows and wooded hill-
sides, sometimes a considerable distance from a stream
(Smith & al., 1993).
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
807
1b. Hymenocallis occidentalis var. eulae (Shinners) G.
Lom. Sm. & Flory, Novon 11: 230. 2001. º
Hymenocallis eulae Shinners, Field & Lab. 19: 103.
1951. – Type: U.S.A. Texas, Van Zandt County: N of
Edgewood near Ocean Lake cultivated at J. A. White
house having been introduced from wild plants on
Sabine River, 6 Sep 1946, Eula Whitehouse 16448
(holotype: BRIT!, isotype: MO!).
Description: Leaves appearing in late winter, with-
ering in late spring before anthesis, suberect, up to 5 cm
wide.
This taxon, originally collected from the banks of the
Sabine River in eastern Texas, is distributed in scattered
sites in the piney woods of Texas, especially along
stream and seepage slopes, and extends northward into
eastern Oklahoma (Correll & Johnston, 1970).
Howard (1995) argued that Hymenocallis galvesto-
nensis (Herb.) Baker was the correct name for this taxon.
See below under H. liriosme (Raf.) Shinners for an
explanation of why we believe that Howard’s argument
is incorrect.
2. Hymenocallis coronaria (Leconte) Kunth, Enum. Pl.
5: 855. 1850. ºPancratium coronarium Leconte,
Ann. Lyceum Nat. Hist. New York 3: 145, pl. 4, fig.
7–9. 1836. ºTomodon coronarium (Leconte) Raf.,
Fl. Tellur. 4: 22. 1838. – Original locality: U.S.A.:
“in Savannah River, at the rapids, a few miles above
Augusta”; no type specimen found. – Lectotype
(designated here): [icon] Leconte, fig. 9. – Epitype
(designated here): U.S.A. Georgia, Lincoln County:
in shallow water at lower end of Price Island in
Savannah River in SE part of county, ca 240 ft, 29
Jun 1949, W. H. Duncan 9840 (GA!).
Description: Bulb nonrhizomatous, globose. Leaves
shiny green, coriaceous, erect but becoming expanded
and reflexed in distal 1/3, strap-shaped, (3.8–) 5–9.5 dm
× 1.5–4.5 cm. Flowers 3–7 or 8–12 per scape, fragrant;
perianth rays white, ascending, 6.5–10 cm × 5.5–9 mm;
staminal cup white with prominent yellow-green eye,
broadly funnelform to chalicelike, 4.5–7 × 5–8.5 cm,
margin often with 2 prominent dentations between the
free portions of the filaments.
Diagnosis: Hymenocallis coronaria is the only
Hymenocallis species with a large, broadly fun-
nelform/chalicelike staminal cup possessing a prominent
yellow-green eye at the base of the cup.
This species inhabits the rocky shoals of Piedmont
river systems above and at the Fall Line in Alabama,
Georgia and South Carolina. Populations at Hargrove
Shoals, Cahaba River, Alabama and the shoals of the
Catawba River, Landsford Canal State Park, South
Carolina may be locally abundant, but the plants do not
occur away from these shoals. The environments where
populations still exist should be protected to ensure the
continued existence of this species (Davenport, 1990a, b,
1996; Smith & al., 1990).
3. Hymenocallis liriosme (Raf.) Shinners, Field & Lab.
19: 102. 1951. ºPancratium liriosme Raf., Fl.
Ludov. 19. 1817. – Original locality: U.S.A.: “moist
grounds and round the small lakes”; no type speci-
men found. – Neotype (designated here): U.S.A.
Louisiana, Terrebonne Parish: from plant in boggy
land near Houma, 15 Apr 1914, J. K. Small 291
(LSU!).
=Hymenocallis galvestonensis (Herb.) Baker, Handb.
Amaryll. 126. 1888. ºChoretis galvestonensis
Herb., Amaryllidaceae 221–222, pl. 41, figs. 34–36.
1837. – Type: U.S.A. Texas: Galveston Bay, 1835,
Drummond III. n. 412. – Lectotype (designated
here): K (“Herbarium Hookerianum”)!; isolecto-
types: BM!, K (“Herbarium Benthamianum”)!
=Hymenocallis galvestonensis subsp. angustifolia
Traub, Pl. Life 23: 67. 1967. – Type: U.S.A.
Alabama, Mobile County: Mobile, 24 Apr 1899,
Biltmore Herbarium H/60 (holotype: US 980051!).
Description: Bulb nonrhizomatous, subglobose to
ovoid. Leaves shiny green, coriaceous, suberect, strap-
shaped to narrowly lanceolate, channelled, 3.2–8.5
(–10.6) dm × 1.7–3.6 cm. Flowers (3–)5–12 per scape,
sweetly fragrant; perianth rays white, slightly ascending,
up to 9 cm × 4–8 mm; staminal cup with prominent yel-
low-green eye, becoming rotate, 2–3.5 × 4–5 cm, with
margin undulate to coarsely dentate between free por-
tions of the filaments.
Diagnosis: Hymenocallis liriosme is similar to H.
coronaria in its staminal cup with a prominent basal yel-
low-green eye, but differs in that the cup is distinctly
smaller, has a coarsely dentate margin and becomes
rotate.
This species inhabits ditches, margins of marshes
and ponds and alluvial woods from the west side of
Mobile Bay to eastern Texas and north to Oklahoma and
Arkansas (Gooch & Smith, 1994; Smith & Coile, 1999).
Howard (1995) argued that Drummond 412, the type
of Hymenocallis galvestonensis (Herb.) Baker, is a spec-
imen of H. eulae Shinners (= H. occidentalis var. eulae)
and therefore H. galvestonensis is the correct name for
H. eulae. For a number of reasons, Smith & Flory (2001)
disagreed. Firstly, the illustration (plate 41, fig. 34)
accompanying Herbert’s (1837) original description of
Choretis galvestonensis, an illustration based on
Drummond 412, shows an inflorescence that is a clear
match for that of H. liriosme. Secondly, the senior author
has examined specimens of Drummond 412 (BM, K) that
without doubt correspond to H. liriosme. Thirdly,
Drummond 412 was collected near Galveston Bay, an
Smith & Garland Nomenclature of Hymenocallis 52 November 2003: 805–817
808
area with numerous wetlands, prime habitats for H.
liriosme. In fact, all collections of Hymenocallis from
Galveston County examined are of H. liriosme; none are
of H. eulae, which prefers a drier habitat. This taxonom-
ic decision preserves the current common usage of the
name H. eulae (Shinners, 1951; Correll & Johnston,
1970; Correll & Correll, 1972).
Hymenocallis galvestonensis subsp. angustifolia
Traub is a narrow-leaved form of H. liriosme that does
not have a definite geographic range. For this reason, we
believe it is not worthy of taxonomic recognition.
4. Hymenocallis choctawensis Traub, Pl. Life 18: 70.
1962. – Type: U.S.A. Florida, Walton County: from
bulbs collected by Mrs. Mary G. Henry, H. P. Traub
263a and b(holotype: MO! [ex TRA]). Fig. 1.
Description: Bulb rhizomatous, ovoid to subglo-
bose. Leaves shiny green, coriaceous, arching to
suberect, channelled proximally, oblanceolate, 3.5–7.5
(–8.5) dm × 2.5–6 cm. Flowers 2–8(–12) per scape, high-
ly fragrant; perianth rays white, slightly ascending to
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
Fig. 1. Hymenocallis choctawensis Traub to illustrate features of the genus. A, habit; B, leaf cross-section (proximal);
C, leaf cross-section (distal); D, section of staminal cup; E, developing fruits; F, seed. Drawn by Mark A. Garland from
Smith & Garland 1729 (Florida, Calhoun Co.: floodplain of Apalachicola River, 12 Jun 1998, HPU).
809
spreading, 8–12 cm × 5–9 mm; staminal cup funnelform,
2.5–4(4.6) × 4–5.5 cm, gradually spreading, margin
irregularly 2- or 3-dentate between the free portions of
the filaments.
Diagnosis: Hymenocallis choctawensis is the only
species of the southeastern United States with shiny,
coriaceous, distinctly oblanceolate leaves, flowers with
noticeably long-spreading white perianth rays and scape
bracts that are triangular and with apex not long-acumi-
nate.
This species occurs from the Apalachicola River
drainage in the central Florida panhandle to south-central
Louisiana and northeastward into southwest Georgia
along the Flint River. Sometimes occurring sympatrical-
ly with Hymenocallis liriosme, it can be distinguished
from that species by a late spring flowering time, by its
distinctly oblanceolate leaves and by its flowers with
longer, widely spreading perianth rays (Smith & al.,
1991, 1996; Smith & Garland, 1996).
5. Hymenocallis crassifolia Herb., Appendix: 44.
1821. ºPancratium crassifolium (Herb.) Schultes,
Syst. Veg. (ed. 16) 7(2): 921. 1830. Original locality:
“Florida” (“ex portu St. Mary, Floridae Orientalis
lat. circit. 29”, Herbert, 1837: 215). – Neotype (des-
ignated here): U.S.A. Georgia, Camden County:
tidal shores of Satilla River, 24 May 1969, W. H.
Duncan 23312 (GA!).
=Pancratium rotatum var. biflorum Ker Gawl., Bot.
Mag. 27: pl. 1082. 1808, as “(a) biflora” º
Hymenocallis mexicana Herb., Appendix: 44. 1821.
Original locality: “the produce of a bulb received by
[Mr. Loddiges] from the same quarter [America]”;
no type specimen found. – Lectotype (designated
here): [icon] Bot. Mag. pl. 1082. 1808.
=Hymenocallis palusvirensis Traub, Pl. Life 18: 71.
1962. – Type: U.S.A. North Carolina, Brunswick
County: near Supply, from bulbs collected by Mrs.
Mary G. Henry (T-151), H. P. Traub 251a,band c
(holotype: MO! [ex TRA]).
Description: Bulb rhizomatous, narrowly ovoid.
Leaves shiny-green, coriaceous, suberect, channelled,
strap-shaped, (2.4–)3.5–6.8 dm × 1–2.5 cm. Flowers 2 or
3 per scape, fragrant; perianth rays white, slightly
ascending, (6–)7–9(–11.5) cm × 3–6 mm; staminal cup
white, funnelform at peak anthesis, gradually spreading,
(2–)2.5–3.5 × 3.4–4.5 cm, with margin between the free
portions of the filaments irregularly dentate, with projec-
tions not prominent.
Diagnosis: Hymenocallis crassifolia is distinguished
by leaves that are shiny, coriaceous, suberect, greater
than 1 cm in width and flowers with a funnelform, sta-
minal cup and perianth rays up to 11.5 cm long.
This species is distributed in ditches, on margins of
streams and rivers, in bogs, brackish marshes and allu-
vial woods from the outer Coastal Plain of the Carolinas
to northeast Florida (Radford & al., 1968; Smith & al.,
1999).
Given the common epithet, Hymenocallis mexicana
Herb. might appear to be a new combination based on
Pancratium mexicanum L. (see “Excluded Taxa and
Names” below). However, Herbert (1821), in his original
publication of the name, did not refer to P. mexicanum or
make any mention of Linnaeus in the relevant portions of
the “Appendix”, and did not provide any descriptive
material, citing only “Bot. Mag. 1082; P. Rotatum biflo-
rum”, referring to P. rotatum var. biflorum Ker Gawl., a
taxon from which Ker Gawler (1808) explicitly excludes
P. mexicanum L. Hymenocallis mexicana Herb. is, there-
fore, technically a new name at species rank for P. rota-
tum var. biflorum. Sealy (1954) and Bauml (1979) also
concluded that Herbert had coined a new name not based
on Linnaeus’ Pancratium mexicanum.
We have determined that Pancratium rotatum var.
biflorum Ker Gawl., the taxon illustrated in plate 1082, is
a synonym of Hymenocallis crassifolia Herb. Plate 1082
shows an Hymenocallis with coriaceous, suberect leaves
and an inflorescence of 2 flowers, each having a small,
funnelform staminal cup at peak anthesis. We associate
these characteristics with H. crassifolia. Because
Herbert’s H. mexicana is based on P. rotatum var. biflo-
rum, it is also a synonym of H. crassifolia.
6. Hymenocallis pygmaea Traub, Pl. Life 18: 70–71.
1962. – Type: U.S.A. South Carolina, Horry County:
small tributary of the Waccamaw River, from bulbs
collected by Mrs. Mary G. Henry (T-129), H. P.
Traub 279 (holotype: MO! [ex TRA]).
Description: Bulb rhizomatous, narrowly ovoid.
Leaves shiny-green, coriaceous, erect, channelled, linear
to narrowly strap-shaped, 1.5–4 dm × 0.5–1.3(–2) cm.
Flowers 1 or 2 per scape, rarely 3, with lemony fra-
grance; perianth rays white, slightly ascending, 5–7 cm ×
3–6 mm; staminal cup white, funnelform at peak anthe-
sis, gradually spreading, 2–3 × ca. 3 cm, with margin
between the free portions of the filaments irregularly
dentate, with projections comparatively small.
Diagnosis: Hymenocallis pygmaea has shiny-green,
coriaceous, erect, narrow leaves and small flowers with
perianth rays not exceeding 7 cm long.
This species, as we interpret it, is endemic to the
Waccamaw River drainage basin in North Carolina and
South Carolina.
7. Hymenocallis duvalensis Traub, Pl. Life 23: 66. 1967.
– Type: U.S.A. Florida, Alachua County: Fairbanks,
swamp, 29 Apr 1928, G. F. Weber s.n. (holotype:
FLAS 6296!; isotype: FLAS 6297!).
Smith & Garland Nomenclature of Hymenocallis 52 November 2003: 805–817
810
Description: Bulb rhizomatous, narrowly ovoid.
Leaves deep-green, low-arching, often appearing pros-
trate, strap-shaped or narrowly oblanceolate, 2.4–5.2 dm
× 1–2.5 cm. Flowers 2 or 3 per scape, rarely more,
intensely fragrant; perianth rays white, slightly ascend-
ing, 7–10 cm × 4–6 mm; staminal cup white, becoming
rotate at peak anthesis, gradually spreading, (2.5–)3–4 ×
4–5 cm, with margin between the free portions of the fil-
aments irregularly and coarsely dentate.
Diagnosis: Hymenocallis duvalensis is the only
Hymenocallis species in the southeastern United States
with strap-shaped to narrowly oblanceolate, arching
leaves and with flowers rotate at peak anthesis but much
smaller than those of H. rotata.
This species occurs along floodplains and stream
banks in Florida, from the Ochlockonee River to north of
Gainesville, and into south-central Georgia along the
Ochlockonee River and its tributaries (Puffenbarger &
Smith, 1993; Smith & al., 1999).
8. Hymenocallis franklinensis G. Lom. Sm., L. C.
Anderson & Flory, Novon 11: 235. 2001. – Type:
U.S.A. Florida, Franklin County: Cow Creek near its
confluence with the Ochlockonee River, 27 Apr
1991, L. C. Anderson 13382 (holotype, FSU!).
Description: Bulb rhizomatous, narrowly ovoid.
Leaves lustrous, coriaceous, nearly erect, channelled,
strap-shaped, 2.8–6 dm × 1.8–2.7 cm. Flowers 2 or 3 per
scape, moderately fragrant; perianth rays white, slightly
ascending, 8.8–11.5 cm × 5–7 mm; staminal cup white,
funnelform at peak anthesis, gradually spreading, 3–4 ×
4–5.5 cm, with margin between the free portions of the
filaments with irregularly dentate projections.
Diagnosis: Hymenocallis franklinensis is distin-
guished by lustrous, coriaceous, suberect leaves 1.8–2.7
cm broad and flowers that are larger in all dimensions
than those of Hymenocallis crassifolia.
This species occurs along the banks of streams in the
lower Ochlockonee River drainage basin in Franklin
County, Florida. Similar plants grow in the adjacent
Sopchoppy River drainage basin. Further investigation is
needed to determine whether these plants belong to this
species.
9. Hymenocallis rotata (Ker Gawl.) Herb., Appendix:
44. 1821. – Pancratium rotatum Ker Gawl., Bot.
Mag. 21: pl. 827. 1805. ºPancratium rotatum var.
pluriflorum Ker Gawl., Bot. Mag. 25: pl. 1082.
1808, as “(b)pluriflora” ºHymenocallis rotata var.
quadrifolia Herb., Amaryllidaceae 217. 1837. º
Tomodon rotatum (Ker Gawl.) Raf., Fl. Tellur. 4: 22.
1838. Original locality: U.S.A. “Carolina”; no type
specimen located. – Lectotype (designated here):
[icon] Bot. Mag. 21: pl. 827. 1805. – Epitype (desig-
nated here): U.S.A. Florida, Wakulla County: mar-
gins of the Wakulla River, near bridge at US 98, 26
May 1990, R. K. Godfrey 83870 (FSU!).
=Hymenocallis disciformis (DC.) M. Roem., Fam.
Nat. Syn. Monogr. 4: 174. 1847. ºPancratium dis-
ciforme DC. in Redouté, Liliac. 3: t. 155. 1806. º
Hymenocallis paludosa Salisb., Trans. Hort. Soc.
London 1: 338. 1812. ºHymenocallis rotata var. dis-
ciformis (DC.) Herb., Amaryllidaceae 217. 1837. –
Lectotype (designated here): [icon] Redouté, Liliac.
3: t. 155. 1806.
=Tomodon floridanum Raf., Fl. Tellur. 4: 22. 1838. º
Hymenocallis floridana (Raf.) C.V. Morton, Year
Book Amer. Amaryllis Soc. 2: 80–83. 1935. – Lecto-
type (designated here): [icon] Leconte, Ann. Lyceum
Nat. Hist. New York 3: 144. pl. 4, fig. 6. 1836.
=Hymenocallis lacera Salisb. var. minor Chapman,
Fl. South. U.S. ed. 3: 494. 1897. ºPancratium rota-
tum b. Leconte, Ann. Lyceum Nat. Hist. New York 3:
144. 1836. – Type: not known.
= Hymenocallis laciniata Small, Man. S.E. Fl. 323,
1503. 1933. – Type: U.S.A. Florida, Taylor County:
Carbur, 6 May 1925, J. K. Small s.n. (holotype: NY;
type specimen not found).
=Hymenocallis floridana subsp. amplifolia Traub, Pl.
Life 23: 67. 1967. – Type: U.S.A. Florida, Marion
County: Oklawaha River at Eureka, 11 May 1935, F.
West, A. N. Tissot & W. B. Tisdale s.n. (holotype:
FLAS 10473!; isotype: FLAS 10474!)
Ker Gawler’s diagnosis and discussion of P. rotatum
var. pluriflorum indicate that he was equating this vari-
ety, and not his alpha variety, P. rotatum var. biflorum
(see H. crassifolium above), with his earlier P. rotatum
Ker Gawl. (Ker Gawler, 1805). Rafinesque based
Tomodon floridanum on the Pancratium rotatum pub-
lished by Leconte in 1836. Leconte made clear, both
from his introduction and from the lay-out of his two
undoubtedly new species (P. coronarium and P. occiden-
tale) that P. rotatum was an existing species, not a new
one, and so he was apparently adopting P. rotatum Ker
Gawl., though he did not cite an author. Rafinesque,
however, specifically excluded P. rotatum Ker Gawl.,
treating it as T. rotatum (Ker Gawl.) Raf. Tomodon flori-
danum is therefore nomenclaturally distinct from P. rota-
tum, to be typified from Leconte’s material, which we
consider to be conspecific with H. rotata (Ker Gawl.)
Herb. For further discussion of H. paludosa Salisb. see
under “Excluded Taxa and Names” below.
Description: Bulb rhizomatous, ovoid. Leaves deep
green, coriaceous, arching to suberect, strongly channel-
led proximally, strap-shaped, 4–10 dm × 2–5 (–7) cm.
Flowers 2–4 per scape, fragrant; perianth rays white,
extending nearly horizontally from base of staminal cup,
9–14.5 cm × 7–15 mm; staminal cup white, rotate at peak
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
811
anthesis, 4–6 × 6–9 cm, with margin often with 2 or 3
prominent lacerate dentations between the free portions
of the filaments.
Diagnosis: Hymenocallis rotata is the only
Hymenocallis species of the southeastern United States
having broad, rotate staminal cups and perianth rays
more than 9 mm wide.
This species occurs along spring runs and adjacent
wetlands south of Tallahassee eastward and southward in
the Florida peninsula (Smith & Darst, 1994).
10. Hymenocallis tridentata Small, Man. S.E. Fl. 323,
1503. 1933. Type: U.S.A. Florida, Indian River
County: Vero, Dec 1920, J. K. Small s.n. (holotype:
NY!).
=Hymenocallis traubii Moldenke, Pl. Life 23: 61.
1967. – Type: U.S.A. Florida, Volusia County:
grown from bulbs collected by E. L. Brasol near
Daytona Beach, H. P. Traub 1056 (holotype: MO!
[ex TRA]).
Description: Bulb rhizomatous, ovoid. Leaves shiny
green, coriaceous, arching, channelled, strap-shaped,
2–5.5 dm × 1–1.8(–2.5). Flowers rarely 1, usually 2 or 3
per scape, fragrant; perianth rays white, slightly ascend-
ing, 7–11(–12.5) cm × 5–8 mm; staminal cup broadly
funnelform to rotate, 3–4.5 × 4–6 cm, with margin often
with 2 or 3 prominent dentations between the free por-
tions of the filaments.
Diagnosis: Hymenocallis tridentata is similar to H.
rotata but its leaves are narrowly strap-shaped and often
less than 5 dm long and its flowers are smaller.
This species occurs in southern Florida in ditches,
marshes and wet prairies. It also occurs sporadically in
wetlands as far north as eastern central Florida (Herndon,
1987).
11. Hymenocallis godfreyi G. Lom. Sm. & Darst,
Novon 4: 398. 1994. – Type: U.S.A. Florida,
Wakulla County: San Marcos de Apalache, marsh
near confluence of Wakulla and St. Marks Rivers, 16
Mar 1990, R. K. Godfrey 83721 (holotype: FSU; iso-
type: FLAS, MO, NY, USF).
Description: Bulb basally rhizomatous, ovoid.
Leaves yellow-green, coriaceous, suberect, channelled,
strap-shaped, (1.3–)2–3.8 dm × 1–2.5 cm. Flowers 2,
rarely 3, per scape, fragrant; perianth rays white, slightly
ascending, 7–10.5 cm × 5–8 mm; staminal cup broadly
funnelform, 3–4.5 × 5 cm, with margin between the free
portions of the filaments often with 2 prominent lacerate
projections.
Diagnosis: Hymenocallis godfreyi is similar to H.
rotata and H. tridentata but is distinguished by a rhizome
that arises from the bottom of the basal plate instead of
laterally, by leaves that are yellowish-green and often no
more than 2–3 dm in length at anthesis.
This is one of the rarest spider-lilies known, occur-
ring only in brackish marshes near St. Marks, Florida.
12. Hymenocallis puntagordensis Traub, Pl. Life 18: 71.
1962. – Type: U.S.A. Florida, Charlotte County:
from bulbs collected by C. L. Burlingham, H. P.
Traub 878a, band c(holotype: MO! [ex TRA])
(Smith & al., 1994; Smith & Garland, 1996).
Description: Bulb nonrhizomatous, narrowly ovoid.
Leaves shiny-green, coriaceous, suberect, channelled
proximally, strap-shaped, 3.5–7.5 dm × 1.5–3 cm.
Flowers 3–5 per scape, fragrant; perianth rays white,
ascending, (7–)9.5–11.5 cm × 4–6 mm; staminal cup fun-
nelform at peak anthesis, 2–2.8 × 2–3 cm, with margin
between the free portions of the filaments with 1 or 2
prominent lacerate projections.
Diagnosis: Hymenocallis puntagordensis is the only
Hymenocallis species in the southeastern United States
with a small funnelform staminal cup having prominent
lacerate projections and with the free portions of the fil-
aments green distally.
This species is known from only a few collections
made near Punta Gorda, Florida. It may be a taxon natu-
ralized from the Neotropics that has undergone natural
selection to develop its unique set of characteristics
(Smith & Garland, 1996).
13. Hymenocallis palmeri S. Watson, Proc. Amer. Acad.
Arts 14: 301. 1879. – Type: U.S.A. Florida, Dade
County: Biscayne Bay in 1874, Edward Palmer 554
(holotype: GH!)
=Hymenocallis humilis S. Watson, Proc. Amer. Acad.
Arts 14: 301. 1879. – Type: U.S.A. Florida: Indian
River in 1874, Edward Palmer 555 (holotype: GH).
See Traub (1956).
Description: Bulb nonrhizomatous, narrowly ovoid.
Leaves blue-green, coriaceous, erect, channelled, linear,
1.5–5(–6.5) dm × 4–10 mm, margin hyaline. Flower 1,
rarely 2, per scape, faintly fragrant; perianth rays yellow-
green to pale green, ascending, 7.5–12.5 cm × 3–7 mm
(unfolded); staminal cup funnelform at peak anthesis,
somewhat spreading, 2.5–3.5 (–5) × 3–5 cm, margin
between the free portions of the filaments irregularly
dentate with some filiform projections.
Diagnosis: Hymenocallis palmeri is the only Hym-
enocallis species of the southeastern United States with a
single flower per scape and green perianth rays.
This species is relatively common in ditches, marsh-
es, fields and wet prairies of central and southern Florida
and is disjunct in northeastern Florida (Smith & Flory,
1990).
14. Hymenocallis henryae Traub, Pl. Life 18: 71: 1962.
Smith & Garland Nomenclature of Hymenocallis 52 November 2003: 805–817
812
– Type: U.S.A. Florida, Walton County: Santa Rosa
Beach, from bulbs collected by Mrs. Mary G. Henry
(T-130), H. P. Traub 282a and b(holotype: MO! [ex
TRA]).
Description: Bulb nonrhizomatous, broadly ovoid.
Leaves blue-green to deep green, coriaceous, erect,
broadly channelled, strap-shaped, (2–)3–6.7 dm ×
(0.8–)1.5–3.2 cm, margin hyaline. Flowers 2, rarely 3,
per scape, mildly fragrant; perianth rays pale green,
slightly ascending to widely-spreading, 8.8–16 cm ×
4.5–9.5(–11) mm; staminal cup funnelform, gradually
spreading, 3–4 × 5–6 cm, with margin between the free
portions of the filaments irregularly dentate (Smith &
Flory, 1990).
Diagnosis: Hymenocallis henryae is similar to H.
palmeri in its green perianth rays but differs in its 2
(rarely 3) large flowers per scape.
14a. Hymenocallis henryae var. henryae
Description: Plants growing singly or in loose, scat-
tere clumps. Leaves light blue-green and slightly glau-
cous to deep shiny green and nonglaucous, (2–) 3–6.7
dm × 0.8–3.2 cm. Perianth rays up to 13 cm long.
This variety occurs in cypress drainages and
stringers associated with wet pine flatwoods in Gulf, Bay
and Walton Counties, Florida.
14b. Hymenocallis henryae var. glaucifolia J.N. Henry
& G. Lom. Sm., Herbertia 54: 114. 2000 (“1999”). –
Holotype: U.S.A. Florida, Liberty County: NW of
Sumatra, 10 Jun 1986, Godfrey 82044 (FSU!).
= “Hymenocallis viridiflora” Small, Man. S.E. Fl. 322,
1933, non rite publ. (Art. 34.1 (a)), non H. viridiflo-
ra (Ruiz & Pav.) R. W. Wallace, 1928.
Small (1933: 322) provided five keys in his treat-
ment of Hymenocallis: a key to four infrageneric groups
and keys to the species within each group. Group III,
called “Viridiflorae” in the key to infrageneric groups
and “Viridiflora” in the key to species, included only
species 10, H. viridiflora. Small’s key (and his text)
described his species 10 as having green petals and se-
pals and two flowers; among United States species of
Hymenocallis, only H. henryae has this combination of
characters. Small, however, treated species 10 in the text
under the name “H. rotatum LeConte” (Small, 1933:
324), and included only that name in the Index (Small,
1933: 1530). It would appear, therefore, that Small did
not accept the name H. viridiflora in the Manual, and,
therefore, it is not validly published under Art. 34.1(a) of
the ICBN. Even if Small had applied the name H. viridi-
flora to his description of species 10, as he may have
originally intended, the name would have been illegiti-
mate since it would have been a later homonym of H.
viridiflora (Ruiz & Pav.) R.W. Wallace, applied to a
Peruvian species. Why Small applied the name “H. rota-
tum LeConte” (Pancratium rotatum Ker Gawler) to his
species is unclear. Leconte describes P. rotatum as hav-
ing eight leaves, broadest at the middle, and four flowers,
while Small describes plants with two or three glaucous
linear leaves and usually two flowers. Small’s inclusion
of “H. viridiflora” in a unispecific group strongly sug-
gests that he considered it highly distinct from other
Hymenocallis. Small’s description of “H. rotatum
LeConte” refers to the leaves as glaucous, the chief dis-
tinguishing feature of H. henryae var. glaucifolia, and so
we conclude that it is to this taxon that Small’s species
no. 10 refers.
Description: Plants growing in dense clumps.
Leaves blue-green from heavy glaucescence, 3–6 dm ×
1.9–2.9 cm. Perianth rays up to 16 cm long.
This variety occurs in cypress depressions at the
edge of pine flatwoods and around cypress domes in the
Apalachicola National Forest, Liberty County, Florida.
15. Hymenocallis latifolia (Mill.) M. Roem., Fam. Nat.
Syn. Monog. 4: 168. 1847. ºPancratium latifolium
Mill., Gard. Dict. ed. 8. Pancratium no 8. 1768. –
Type: Jamaica: ports of St. Christopher and
Barbados, H. Sloane s.n. (Herb. IV. 112) (holotype:
BM! as seen on web site)
=Hymenocallis caymanensis Herb., Amaryllidaceae
214. 1837. – Original locality: “Grand Cayman”;
type specimen not found.
=Hymenocallis collieri Small, Man. S.E. Fl. 322-323,
1503. 1933. – Type: U.S.A. Florida, Collier County:
Marco Island, May 1920, J. K. Small s.n. (holotype:
NY; type specimen not located).
=Hymenocallis keyensis Small, Man. S.E. Fl. 322.
1933. – Based on Hymenocallis caribaea sensu
Chapman, Fl. South. U.S. ed 2: 654. 1883, non (L.)
Herbert. – Type: not known (original locality:
[U.S.A. Florida] “mangrove swamps, sandy shores,
and rarely hammocks, eastern coast of pen. Fla. and
the Keys”).
=Hymenocallis kimballiae Small, Man. S.E. Fl. 323,
1503. 1933. – Type: U.S.A. Florida, Franklin
County: Apalachicola, May 1921, Winifred Kimball
s.n. (holotype: NY; type specimen not found).
Description: Bulb nonrhizomatous, broadly ovoid,
forming numerous bulblets. Leaves lustrous, coriaceous,
arching and spreading distally, broadly liguliform, 5–10
dm × 4–9 cm. Flowers 9–15 per scape, sweetly fragrant:
perianth rays white, extending nearly horizontally,
reflexed distally, 7.5–13 cm × 4–6 mm; staminal cup
funnelform, 2–3 × 3–4 cm, margin between the free por-
tions of the filaments wavy, without prominent projec-
tions.
Diagnosis: Hymenocallis latifolia is the only
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
813
Hymenocallis species of the southeastern United States
with broadly strap-shaped leaves and orange, rather than
yellow, pollen.
This species inhabits coastal dunes and swales, sand
ridges, edges of flatwoods and mangrove swamps,
chiefly in southern Florida but it also occurs sporadical-
ly north along the coast to the panhandle.
EXCLUDED TAXA AND NAMES
Hymenocallis caroliniana (L.) Herb., Appendix: 44.
1821. ºPancratium carolinianum L., Sp. Pl. 291. 1753.
Original locality: “Jamaica, Carolina”; no type specimen
found. – Lectotype (designated here): [icon] Catesby,
Nat. Hist. Carolina Appendix: fig. 5. 1747.
Linnaeus based Pancratium carolinianum entirely
on Catesby’s illustration and description of Lilio-
Narcissus Polianthus, flore albo in the Appendix to his
Natural History of Carolina (figure 5, page 5); published
in 1747 (Stafleu & Cowan, 1976). Catesby said he found
the plant “in a bog near Pallachucula, an Indian Town on
the Savanna river within the precinct of Georgia.”
Pallachucula, Pallachucola or Palichucola appears as the
name of a lake, river bluff and school on older maps of
the area around Brighton, near the Savannah River in
southern Hampton County, South Carolina. It is unclear
why Linnaeus listed Jamaica as an additional locality for
this species. Perhaps the “yellow and black pye”, a trou-
pial, Icterus icterus (Linn.), drawn perching on the
exposed bulb (Fig. 2) and reported by Catesby as
Jamaican, misled Linnaeus into believing that the plant
was also Jamaican.
Catesby’s description is consistent with features of
Hymenocallis, but is too general to determine the species
concerned. At Pallachucula, he may have seen
Hymenocallis crassifolia Herb.
Catesby’s illustration (Fig. 2), on the other hand,
clearly depicts the Mediterranean Pancratium mariti-
mum L., with numerous, narrowly strap-shaped, some-
times twisted leaves, a short scape bearing a several-
flowered inflorescence, each flower with relatively short
and broad tepals, a large campanulate staminal cup with
2 triangular lobes between each pair of stamens, and
short filaments. Ker Gawler (1808) said of the plate, “we
have very little doubt but that it is in reality the repre-
sentation of the European maritimum, which Catesby
probably saw in bloom at some of the English gardens of
that day; when judging from general appearance, he
believed it to be the same with the plant which he says he
had seen in America...”. Fernald & Schubert (1948),
however, claimed that Catesby’s plate was a “beautiful”
representation of Hymenocallis coronaria (Leconte)
Kunth, and Sealy (1954) believed that a collection from
Alabama represented at BM and K (A. H. Howell s. n. in
1913), which the senior author here has identified as H.
coronaria, matched Catesby’s figure and description. We
disagree. Hymenocallis coronaria has long coriaceous
leaves, erect for two-thirds of their length, that are not
twisted, tall scapes and irregular teeth on the margin of
the staminal cups, not the broad, regular triangular teeth
as depicted in Catesby’s plate; moreover, Catesby’s
locality of occurrence for the species, a bog near
Pallachucula, is in the swampy lower Coastal Plain near-
ly 130 km downriver from the rocky shoals in the
Piedmont where H. coronaria is known to grow.
Catesby’s plate and description are the only elements in
the Linnaean protologue. By choosing Catesby’s figure 5
as the lectotype of Pancratium carolinianum, we make
this name a synonym of P. maritimum L., Sp. Pl. 291.
1753, and hence exclude it from the scope of this treat-
ment.
Hymenocallis lacera Salisb., Trans. Hort. Soc.
London 1: 338. 1812, nomen superfluum et illegitimum.
Salisbury gives no description and lists two syn-
onyms: Pancratium rotatum [var.] bKer Gawler and
Smith & Garland Nomenclature of Hymenocallis 52 November 2003: 805–817
814
Fig. 2. Figure 5 in the “Appendix” to Mark Catesby’s
Natural History of Carolina, Florida, and the Bahama
Islands (1747), showing Lilio-Narcissus Polianthus, flore
albo, the lect ot yp e of Pancratium carolinianum L.
Copyright © Board of Regents of the University of
Wisconsin System. All rights reserved.
Pancratium carolinianum L. Without further description,
the species to which Salisbury intended this name to
apply must remain uncertain. However, as Salisbury does
not exclude the type of P. carolinianum, his name is
superfluous and illegitimate and, as he does not cite a
type, is to be typified by the type of Linnaeus’ name;
hence H. lacera Salisb. must also refer to P. maritimum.
Hymenocallis paludosa Salisb., Trans. Hort. Soc.
London 1: 338. 1812, nomen superfluum et illegitimum.
Salisbury gives no description and lists three syn-
onyms: Pancratium rotatum [var.] a“Ker Gawler,
Pancratium disciforme DC., and Pancratium mexicanum
as used by Michaux in Flora Boreali-Americana 1: 188.
1803. As each of these three synonyms could refer to a
different species, it is not clear what taxon Salisbury had
in mind. However, we have been able to assign
Pancratium disciforme DC. as a synonym of Hymen-
ocallis rotata (Ker Gawl.) Herb. As Salisbury cites P. dis-
ciforme DC. without any exclusion of its type, the name
H. paludosa is a superfluous and illegitimate name,
homotypic with H. disciformis.
“Pancratium fluitans W. Bartram”, Travels Carolina:
35. 1791, nomen nudum.
In describing the vegetation around Augusta,
Georgia, Bartram (1791, p. 35) mentioned “the odorifer-
ous Pancratium fluitans which almost alone possesses
the little rocky islets” in the Savannah River. It appears
from the habitat that he was describing Hymenocallis
coronaria, but he presented no description or illustration
of the plant.
Pancratium mexicanum L., Sp. Pl. 290. 1753. –
Lectotype (designated by Sealy, 1954, p. 217): [icon in]
Dillenius, Hortus Elthamensis t. 222, f. 289. 1732.
Walter (1788), Michaux (1803), Pursh (1814), Elliott
(1821), among others, referred various southeastern spi-
der-lilies to Pancratium mexicanum. The illustration of
“Pancratium Mexicanum, flore gemello candido” in
Dillenius’s Hortus Elthamensis (t. 222, f. 289), the only
original element in Linnaeus’ protologue, shows charac-
teristics that suggest it is not a taxon of the southeastern
United States. The staminal cup illustrated is small but
the free portions of the filaments are exceedingly long
and each bears an anther that produces orange pollen
(“farina crocea”). Also, the leaves appear relatively short
and are broadly oblanceolate. We conclude from these
characteristics that the illustration represents a
Neotropical taxon.
Recent workers have also considered P. mexicanum
to be neotropical. Sealy (1954) and Bauml (1979) syn-
onymized P. mexicanum with Mexican species of
Hymenocallis not found in the United States, Hymeno-
callis dillenii Roemer (Sealy, 1954) and H. harrisiana
Herb. (Bauml, 1979). Dr. Alan Meerow, some years ago,
also informed the senior author that Dillenius’ plate illus-
trated a Hymenocallis species native to Mexico. We have
therefore excluded Pancratium mexicanum L. as a
species within the study area or its name as a synonym of
any name accepted here.
Hymenocallis mexicana Herb. (1821), on the other
hand, is technically a new name at species rank for P.
rotatum var. biflorum Ker Gawler. See discussion under
H. crassifolia above.
ACKNOWLEDGEMENTS
We acknowledge the late Dr. Walter S. Flory for his excellent
efforts and interactions concerning the cytotaxonomy of
Hymenocallis and the late Dr. Robert K. Godfrey for collecting
numerous excellent Hymenocallis specimens, gathering bulb and
seed collections, and for assistance in the field. We thank Angus
Gholson for valuable field assistance and the opportunity to use his
private herbarium, and C. Craig Moretz, Melanie Darst, Josephine
de N. Henry, Gary Knight, Linda Chafin, Kathy Burks, Catherine
M. Bush, Kevin Preuss and Drs. John D. Tobe, Loran C. Anderson,
Ann F. Johnson, Nancy C. Coile, John B. Nelson, Bert Pittman,
Richard Carter, Larry J. Davenport and Fred T. Yeats for valuable
field assistance. We thank numerous botanists and individuals at
the following agencies/institutions for their keen assistance:
Florida Department of Environmental Protection, Florida Natural
Areas Inventory, Florida State University, Henry Foundation for
Botanical Research, High Point University and the United States
Geological Survey. Melanie Darst and Jean P. Hancock are
thanked for being excellent illustrators of Hymenocallis and for
their enthusiasm for the genus. Dr. Wendy Zomlefer is thanked for
technical assistance and providing valuable field collections.The
following herbaria made loans of their Hymenocallisspecimens or
granted permission for visits for their study: BM, BRIT, CLEMS,
F, FAU, FLAS, FSU, FTG, GA, GH, IBE, K, LSU, MISS, MO, P,
PH, NCU, NY, TAMU, TEX (including LL), UNA, UNCC, US,
USCH, USF, and VSC. Dr. Nancy Morin, as convening editor for
Flora of North America, is thanked for facilitating a trip to MO for
the study of the Traub Herbarium. Drs. Pat Holmgren and Jackie
Kallunki are thanked for diligently searching for NY Hymenocallis
specimens of Dr. J. K. Small and making a loan of them to HPU.
Dr. Michael Fay, Royal Botanic Gardens Kew, is thanked for dis-
tinguishing the type specimens of Hymenocallis galvestonensis
,
and Kent Perkins, FLAS, for supplying the numbers of the type
specimens of Hymenocallis duvalensis
. The Libraries of the
University of Wisconsin-Madison graciously allowed us to repro-
duce their digital version of Figure 5 from Catesby’s Appendix as
our Figure 2. We thank Dr. Alan Meerow for editorial reviews and
helpful comments about Hymenocallis. Dr. John McNeill,
Nomenclature Editor, Taxon is extended special gratitude for his
diligence with our manuscript and for his patience and guidance in
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
815
clarifying a number of complex nomenclatural issues. Dr. Kanchi
Ghandhi and two anonymous reviewers are thanked for their valu-
able insights and comments on our paper. Field and laboratory
research undertaken for this paper was funded in part by a North
Carolina Board of Science and Technology grant (no. 90SE010)
and High Point University.
LITERATURE CITED
Baker, J. G. 1888. Handbook of the Amaryllideae. George Bell
& Sons, London. [pp. 120–129.]
Bartram, W. 1791. Travels Through North and South
Carolina, Georgia, East & West Florida. James &
Johnson, Philadelphia.
Bauml, J. W. 1979. A Study of the Genus Hymenocallis
(Amaryllidaceae) in Mexico. M.S. thesis, Cornell Univ.,
Ithaca, N.Y.
Correll, D. S. & Correll, H. B. 1972. Aquatic and Wetland
Plants of the Southwestern United States. Environmental
Protection Agency, Washington, D. C. [Pp. 665–667.]
Correll, D. S. & Johnston, M. C. 1970. Manual of the
Vascular Plants of Texas. Texas Research Foundation,
Renner. [Hymenocallis, pp. 414–415.]
Davenport, L. J. 1990a. Distribution, habitat requirements and
distinct nature of Hymenocallis coronaria. A. S. B. Bull.
37: 56. [Abstr.]
Davenport, L. J. 1990b. The cahaba lily. Alabama Heritage
16: 24–29.
Davenport, L. J. 1996. The cahaba lily: its distribution and
status in Alabama. J. Alabama Acad. Sci. 67: 222–233.
Elliott, S. 1821. A Sketch of the Botany of South-Carolina and
Georgia, vol. 1. J. R. Schenck, Charleston, South
Carolina.
Fernald, M. L. & Schubert, B. G. 1948. Studies of American
types in British herbaria. Rhodora 50: 149–176, 181–208,
217–233.
Flory, W. S. 1976. Distribution, chromosome numbers and
types of various species and taxa of Hymenocallis.
Nucleus (Calcutta) 19: 204–227.
Flory, W. S. 1978. Known distributions of Hymenocallis
Salisbury in North and Middle America and the West
Indies. Pl. Life 34: 47–59.
Godfrey, R. K. & Wooten, J. W. 1979. Aquatic and Wetland
Plants of the Southeastern United States: Monocoty-
ledons. Univ. Georgia Press, Athens. [Pp. 602–603.]
Gooch, R. & Smith, G. L. 1994. Distinctions between
Hymenocallis liriosme (Raf.) Shinners and Hymenocallis
choctawensis Traub. J. Elisha Mitchell Sci. Soc. 110: 25.
[Abstr.]
Greuter, W., Barrie, F. R., Burdet, H. M., Chaloner, W. G.,
Demoulin, V., Hawksworth, D. L., Jørgensen, P. M.,
Nicolson, D. H., Silva, P. C., Trehane, P. & McNeill, J.
1994. International Code of Botanical Nomenclature
(Tokyo Code). Koeltz, Königstein. [Regnum Veg. 131.]
Greuter, W., McNeill, J., Barrie, F. R., Burdet, H. M.,
Demoulin, V., Filgueiras, T. S., Nicolson, D. H., Silva, P.
C., Skog, J. E., Trehane , P., Turland, N. J. &
Hawksworth, D. L. 2 00 0. Inte rn ational Code of
Botanical nomenclature (St. Louis Code). Koeltz,
Königstein. [Regnum Veg. 138.]
Herbert, W. 1821. Appendix [to the Botanical Magazine and
the Botanical Register]. James Ridgeway & Sherwood,
Neely & Jones, London.
Herbert, W. 1837. Amaryllidaceae
. Reprint 1970. J. Cramer,
Lehre, New York.
Herndon, A. 1987. A morphometric comparison of
Hymenocallis palmeri and Hymenocallis floridana
(Amaryllidaceae) in southern Florida. Sida 12: 295–305.
Howard, T. M. 1995. The correct identity of Hymenocallis
galvestonensis (Herb.) Amaryllidaceae
. Herbertia 50:
48–59. [Volume 50 spans two years from 1994 to 1995.
The effective date of publication of this article is 15 June
1995.]
Ker Gawler, J. B. 1805. Pancratium rotatum. Large crowned
Pancratium. Bot. Mag. pl. 827.
Ker Gawler, J. B. 1808. Pancratium rotatum (a). American
Pancratium. Bot. Mag. pl. 1082.
Kunth, K. S. 1850. Enumeratio Plantarum, vol. 5. J. G. Colla,
Stuttgart & Tübingen. [Pp. 664–680.]
Laferriere, J. E. 1996. Validation of names in Hymenocallis
(Amaryllidaceae). Herbertia 51: 66–67.
Leconte, J. E. 1836. Observations on the United States species
of the genus Pancratium. Ann. Lyceum Nat. Hist. New
York 3: 142–148.
Mabberley, D. J. 1997. The Plant-Book, ed. 2. Cambridge
Univ. Press, Cambridge.
Meerow, A. W., Fay, M. F., Chase, M. W., Guy, C. L., Li, Q.-
B., Snijman, D. & Yang, S.-L. 2000a. Phylogeny of
Amaryllidaceae
: molecules and morphology. Pp. 372–386
in: Wilson, K. L. & Morrison, D. A. (eds.), Monocots II:
Systematics and Evolution. Melbourne.
Meerow, A. W., Guy, C. L., Li, Q.-B. & Uang, S.-L. 2000b.
Phylogeny of the American Amaryllidaceae based on
nrDNA ITS sequences. Syst. Bot. 25: 708–726.
Michaux, A. 1803. Flora Boreali-Americana, vol. 1. Levrault
brothers, Paris & Strasbourg.
Morton, C. V. 1935. A chec k list of t he bulbous
Amaryllidaceae native to the United States. Yearbook
Amer. Amaryllis Soc. 2: 80–83.
Puffenbarger, S. & Smith, G. L. 1993. Biosystematic investi-
gations of Hymenocallis crassifolia Herbert and
Hymenocallis duvalensis Traub in the southeastern United
States. J. Elisha Mitchell Sci. Soc. 109: 1. [Abstr.]
Pursh, F. 1814. Flora Americae Septentrionalis
. Printed for
White & Cochrane & Co., London.
Radford, A. E., Ahles, H. E. & Bell, C. R. 1968. Manual of
the Vascular Flora of the Carolinas. Univ. North Carolina
Press, Chapel Hill. [P. 320.]
Roemer, M. K. 1847. I. Amaryllidaceae. -3. Narcissineae.
Syn.
Monogr. 4: 166–177.
Ross, R. 2002. Which illustrations are original material? Taxon
51: 523–524.
Salisbury, R. A. 1812. On the cultivation of rare plants. Trans.
Hort. Soc. London 1: 338–341.
Sealy, J. R. 1954. Review of the genus Hymenocallis. Kew
Bull. 2: 201–240.
Shinners, L. H. 1951. The north Texas sp ec ies of
Hymenocallis (Amaryllidaceae). Field & Lab. 19:
102–104.
Small, J. K. 1933. Manual of the Southeastern Flora.
Published by the author, New York. [Pp. 321–324.]
Smith & Garland Nomenclature of Hymenocallis 52 November 2003: 805–817
816
Smith, G. L. & Coile, N. C. 1999. Distinctions between
Hymenocallis choctawensis and Hymenocallis liriosme.
Assoc. Southeast. Biol. Bull. 46: 242. [Abstr.]
Smith, G. L. & Darst, M. 1994. A n ew species of
Hymenocallis (Amaryllidaceae) in the Florida panhandle.
Novon 4: 396–399.
Smith, G. L. & Flory, W. S. 1990. Studies on Hymenocallis
henryae (Amaryllidaceae) in the Florida panhandle.
Brittonia 42: 212–220.
Smith, G. L. & Flory, W. S. 2001. Hymenocallis
(Amaryllidaceae) in Texas, with a new varietal combina-
tion. Novon 11: 229–232.
Smith, G. L. & Flory, W. S. 2002. Hymenocallis. Pp. 283–293
in: Flora of North America Editorial Committee (ed.),
Flora of North America North of Mexico, vol. 26. Oxford
Univ. Press, New York.
Smith, G. L., Flory, W. S. & Garland, M. A. 1991. Studies on
Hymenocallis choctawensis Traub in the Florida panhan-
dle. Assoc. Southeast. Biol. Bull. 38: 122. [Abstr.]
Smith, G. L., Flory, W. S. & Nelso n, J. B. 1990.
Cytotaxonomic studies on Hymenocallis coronaria (Le
Conte) Kunth in South Carolina Piedmont river systems.
Assoc. Southeast. Biol. Bull. 37: 141. [Abstr.]
Smith, G. L. & Garland, M. A. 1996. Taxonomic status of
Hymenocallis choctawensis and Hymenocallis puntagor-
densis (Amaryllidaceae). Sida 17: 305–319.
Smith, G. L., Garland, M. A. & Flory, W. S. 1994.
Rediscovery of Hymenocallis puntagordensis Tra ub.
Assoc. Southeast. Biol. Bull. 41: 239. [Abstr.]
Smith, G. L., Joye, D. B. & Flory, W. S. 1993. Studies on
Hymenocallis occidentalis (Le Conte) Kunth. Assoc.
Southeast. Biol. Bull. 40: 148. [Abstr.]
Smith, G. L., Moore, M. O. & Flory, W. S. 1996. The spider-
lilies of Georgia. Assoc. Southeast. Biol. Bull. 43: 153.
[Abstr.]
Smith, G. L., Moretz, C. C. & Yeats, F. T. 1999. Spider-lilies
of the Atlantic Coastal Plain. Assoc. Southeast. Biol. Bull.
46: 240. [Abstr.]
Stafleu, F. A. & Cowan, R. S. 1976. Taxonomic Literature:A
Selective Guide to Botanical Publications and Collections
with Dates, Commentaries and Types, ed. 2, pt. 1. Regnum
Veg. 94. Bohn, Scheltema & Holkema, Utrecht.
Traub, H. P. 1956. The holotype of Hymenocallis palmeri.
Taxon 5: 195–196.
Traub, H. P. 1962. Key to the subgenera, alliances and species
of Hymenocallis. Pl. Life 18: 55–72.
Vassar, M. G. & Meerow, A. W. (eds.). 1997. Retraction of
Laferriere, J., 1996, Validation of names in Hymenocallis
(Amaryllidaceae), Herbertia 51: 66–67. Herbertia 52:
10–11.
Walter, T. 1788. Flora Caroliniana. J. Fraser, London.
[Reprint by Murray Publishing Co. for the Arnold
Arboretum, 1946.]
Smith & Garland Nomenclature of Hymenocallis52 November 2003: 805–817
817