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A MONOGRAPH OF NYMPHAEA SUBGENUS HYDROCALLIS
(NYMPHAEACEAE)
John H. Wiersema
Department of Botany
University of Maryland
College Park, Maryland 20742
Abstract. Nymphaea subgenus Hydrocallis is described to include 14 species. Evidence for this
classification was obtained from numerical taxonomy, scanning electron microscopy of seeds and
pollen, chromosome numbers, floral biology, artificial hybridization, flavonoid chemistry, and general
morphology. Representative species from other subgenera of Nymphaea were included in many of
these investigations for comparative purposes. Seed investigations were exceedingly valuable in defin
ing certain species and in clarifying their phylogenetic relationships. Pollen studies, on the contrary,
were of little value in distinguishing individual species but provided evidence for the relationship of
subg. Hydrocallis to other subgenera. The addition of information on chromosome number and
flavonoid profile for 12 taxa, supplemented by observations on floral biology and crossability, has
permitted a general phylogenetic interpretation of subg. Hydrocallis. Two major evolutionary lines are
apparent. One includes N. amazonum, N. prolif?ra, N. lasiophylla, and N. lingulata. Hybridization
events within this line involving subg. Lotos may have contributed to the formation of N. rudgeana,
but more evidence is needed to substantiate this interpretation. The other lineage includes N. novo
granatensis, N. gardneriana, N. conardii, and N. jamesoniana, with a secondary branch involving N.
glandulifera, N. potamophila, N. oxypetala, and perhaps N. belophylla. Nymphaea tenerinervia exhib
its affinities to both major lines and appears to represent an evolutionary link between them. A
complete taxonomic treatment of subg. Hydrocallis, including the description of the new subspecies N.
amazonum subsp. pedersenii, is presented.
INTRODUCTION
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2 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
TAXONOMIC HISTORY
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 3
Seed Morphology. Seeds were obtained in most cases from herbarium speci
mens. Much of this material was the result of my own field collections, although a
small percentage of such seeds was obtained from borrowed herbarium speci
mens. In other instances seeds were obtained from plants in outdoor cultivation,
and in two such instances the seeds were the result of artificial hybridizations.
Available material from 12 species of subg. Hydrocallis was initially scrutinized
with the aid of a dissecting microscope. At that time, measurements of seed
length and width were taken, seed color was noted, and representative collections
were selected for more intensive light and scanning electron microscopy. Seeds of
four additional species of Nymphaea, representing the subgenera Nymphaea, Lo
tos, and Brachyceras, were also included for comparative purposes. Seed material
from 86 populations of these 16 species was initially examined; the 49 collections
receiving more intensive study are listed in Wiersema (1984b).
Three procedures were employed in examining the seed coat morphology of
these seeds. 1) In an SEM study of surface topography, seeds without arils were
required. These were selected from specimens; if such seeds were unavailable, the
aril was mechanically removed after soaking of seeds in Aerosol OT solution.
Seeds were then mounted on aluminum stubs with double adhesive transparent
tape. The samples were coated to 250 ? with a gold/palladium alloy in a Technics
Hummer V Sputter Coater, observed at 20 kV with an ETEC Autoscan SEM,
and photographed with Polaroid Type 55 P/N film. 2) For further examination of
surface cells, the outer testa was detached with forceps from softened seeds. This
testa was mounted in Hoyer's solution on slides, and examined and photographed
with a camera-equipped compound microscope. With the aid of a micrometer, ten
cells from each slide were measured for length, width, and the extent of intercala
tion at midcell. Because cell shape varies considerably over the seed surface, only
cells located medially in regular longitudinal rows were measured to provide a
more standardized estimate of cell shape for each collection of seeds. 3) To aid in
interpreting seed coat anatomy, seeds of all 16 species studied were fixed in 70%
ethanol-formalin-acetic acid, dehydrated in a tertiary butyl alcohol series, embed
ded in Paraplast (melting point, 56-57?C), sectioned at 8-15 jim on a rotary
microtome, and stained with safranin and fast green.
Pollen Morphology. All pollen for scanning electron microscopic analysis was
obtained from herbarium specimens, most of these from my own recent field
collections. The 30 populations furnishing pollen for SEM studies are listed in
Wiersema (1984b). Represented among these were 12 species of subg. Hydrocallis
as well as certain species of subgenera Lotos, Brachyceras, and Nymphaea. Pollen
preparation followed the procedure of Lynch and Webster (1975).
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4 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Chromosome Numbers. All counts were obtained from root tips, mostly using
an aceto-orcein staining procedure following pretreatment with 8-hydroxyquino
line. A total of 25 populations were examined for their chromsome numbers
(Appendix 1). Voucher specimens are deposited at UNA.
Floral Biology and Artificial Hybridization. Although field observations were
made at the time of collection in Mexico, Ecuador, Venezuela, Argentina, Brazil,
and Florida, the majority of observations were garnered from plants cultivated
from field-collected tubers. Plants were cultivated outdoors in Tuscaloosa, Ala
bama (33?N latitude) during May to October of 1982-1984; the majority of data
was gathered during 1983. A total of 28 populations of subg. Hydrocallis were
cultivated though not all progressed to flowering. They were grown in galvanized
washtubs (No. 3 size) in accordance with the recommendations of Pring (1941,
1949).
Several procedures were employed with flowers at anthesis. As no potential
pollinators were ever observed in any of the flowers, the latter were not bagged
but were merely tagged the day of the treatment. Some flowers were allowed to
develop normally in the absence of cross-pollination to test for autogamous or
agamospermous seed production. If seed production resulted from this treatment,
other flowers of the same plant were emasculated, allowed to develop without
pollination, and eventually tested for seed set. Other emasculated flowers were
artificially pollinated with pollen of another available flower (almost always
another species). This established that the effects of emasculation were not the
result of injury to flowers and enabled an assessment of the hybridization poten
tial between various species. To increase the number of available crosses, anthers
of some flowers were removed and allowed to dehisce in sealed plastic containers,
thus allowing these flowers both to serve as a source of pollen and to be polli
nated. Pollen treated in this manner exhibited no apparent loss of viability.
Subsequent to these procedures fruits were allowed to mature, and the seeds
or aborted ovules were collected at the time of dehiscence or decay of the flower
or fruit. Seeds or ovules from a given treatment were spread on p?trie dishes,
oven-dried, and later assayed for the percentages fully developed, partially devel
oped, and aborted. Pollen fertility was also examined for many species with the
use of aceto-carmine jelly according to standard procedures (Radford et al. 1974).
Data on crossing experiments were meant to be interpreted loosely in view of
the lack of replicate crosses and the possible effects of environmental factors, as
each developing fruit experienced a somewhat different set of environmental
conditions. Therefore, rigorous controls on the collection and assessment of seeds
were not enforced. Limitations in the number of potential crosses which became
available precluded a more thorough investigation. Nevertheless, a total of 53
artificial crosses was attempted among members of subg. Hydrocallis, an addi
tional 8 crosses were carried out between individuals of subg. Hydrocallis and N.
lotus of subg. Lotos, and one cross involved N. amazonum and N. odor ata of
subg. Nymphaea.
Flavonoid Chemistry. Leaf material for flavonoid analysis came from: 1) air
dried leaves from field collections, 2) air-dried leaves from mostly outdoor culti
vations, and 3) fresh leaves from outdoor cultivation. The majority of material
was of the first type. Leaves were crushed or fragmented by hand and subjected
to three 24 hour extractions in an aqueous methanol solution which was succ?s
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 5
sively decreased to ca. 50% methanol. The first extraction was carried out under
slight heating; the latter two with agitation. The combined methanol-water eluates
were evaporated under vacuum on a rotary evaporator to an aqueous solution.
Following several extractions of this solution with chloroform, flavonoids were
partitioned into ethyl acetate. The ethyl acetate was then evaporated to dryness
under vacuum, and the flavonoids redissolved in absolute methanol for use in
two-dimensional paper chromatography. Chromatographie separations were
achieved using tertiary butanol:glacial acetic acidiwater (3:1:1 by volume) as one
solvent and 15% aqueous acetic acid as the second solvent on Whatman 3MM
paper, and followed the procedures of Mabry et al. (1970). Further separations of
compounds unresolvable on paper were carried out on 0.1 mm Cellulose MN300
and 0.1 mm Polyamide-TLC6 precoated plates. Best results on polyamide TLC
plates were observed using a solvent system of 1,2-dichloroethane:methanol:
butanone:water (50:25:21:1 by volume).
Purified compounds were eluted from Chromatographie paper or plates with
methanol or, if necessary, compounds were purified using Sephadex LH-20 or
Polyamide SC6 column chromatography. Similarity of compounds from different
populations was detected by cochromatography on TLC precoated plates in a
minimum of three solvent systems (at least one of these on polyamide). Examina
tion of these plates was aided by fuming with ammonia and spraying with Natur
stoffreagenz A.
For identification of flavonoid compounds, the six standard UV spectra were
run in methanol (Mabry et al. 1970). Compounds were then hydrolyzed in boiling
6% HC1 for one hour. The aglycone was recovered in ethyl acetate for UV
spectral analysis and cochromatography with known standards. The aqueous por
tion was subjected to circular chromatography on 0.1 mm cellulose 400 TLC
plates with sugar standards using ethyl acetate:pyridine:water (6:3:2 by volume).
The sugars were visualized with aniline pthalate spray. Rf values of isolated com
pounds were determined on paper in various solvent systems and compared with
published values. For further characterization of certain compounds, proton
NMR spectra were obtained in deuterated DMSO.
A total of 54 populations representing 17 species and several assumed hybrids
were examined for their flavonoid profiles. These included three members of
Nymphaea subg. Nymphaea, two of subg. Brachyceras, and one of subg. Lotos,
with the remainder belonging to subg. Hydrocallis.
To examine the relationships among the various species of subg. Hydrocallis
and to explore their affinities to other members of Nymphaea, a statistical analysis
was applied to the chemical data by using the NT-SYS statistical package (Rohlf
et al. 1977). Two separate data matrices were subjected to cluster analyses. In
both matrices Jaccard coefficients were used to represent the degree of associa
tion between each pair of OTUs. This coefficient is determined by the:
for any two OTUs. The first matrix was computed from data of the 56 compounds
isolated for the 54 populations or OTUs examined. As this analysis was incapable
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6 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Habit. Species of Nymphaea are all perennial aquatic herbs. They are rooted
in what is always a rich organic substrate at the bottom of ponds or sluggish
streams, with the leaves and flowers elevated to the water surface on elongate
petioles and peduncles. A single mature plant may occupy an area from Vi m in
diameter in smaller forms to 3 m or more in larger forms. Flowers are floating or
may be emergent as much as 40 cm on stout peduncles. In Nymphaea subg.
Hydrocallis they are mostly floating, although a few species have slightly emer
gent flowers. Fruits of Nymphaea develop under water, being retracted there by a
bending or coiling of the peduncle; in subg. Hydrocallis an elongated S-shaped
bending is observed in most species.
Plants of Nymphaea withstand seasonal periods of cold or drought either as
seeds or as tuberous rhizomes. Although in subg. Hydrocallis tubers have not
demonstrated an ability to survive freezing temperatures, they are somewhat
resistant to drought. In fact, following cessation of growth at least a brief period
of drying out appears to be necessary for resprouting of tubers. Seeds of these
species may germinate directly following their release or may require a period of
drought.
Stems. The stems of some species of Nymphaea are elongate and horizontal,
but those of subg. Hydrocallis usually consist of an erect caudex a few centimeters
in length, although it may be up to several centimeters long in some species.
Inland species may develop a smooth and globose tuber or one that is more ovoid
and densely woolly. In many species these tubers are developed on lateral shoots,
either at the nodes of elongate stolons or, as in two species, within abortive
tuberiferous flowers. Certain species produce stolons only at the time of germina
tion of tubers; in others stolons are formed throughout the vegetative period and
are an effective means of vegetative propagation.
In species which do not form additional tubers, a tuberous rhizome serves as
the resting state. Several sprouts may develop from this dormant structure at the
onset of the next growing season. The longevity of individual plants which must
perennate in this fashion may be considerably reduced, as such a rhizome in
cultivation seems to remain viable only a few years. All species which fit this
pattern rely heavily on autogamous seed production for their survival and disper
sion. In N. rudgeana, which seems to inhabit more permanent aquatic habitats, a
true resting tuber may not be formed, with the plant maintaining constant growth.
This species also relies extensively on autogamy.
Roots. Although the petiolar and peduncular bases readily decay and absciss
from tubers at the onset of dormancy, certain roots are somewhat more persistent
and may perform a useful function. Roots attached to tubers of several species of
subg. Hydrocallis have been observed to be highly contracted, this contraction
probably serving to retract the tuber well into the substrate as the dry season
progresses. Conard (1905) indicates that contractile roots of other subgenera of
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 7
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8 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 9
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10 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 1. Unusual anatomical features of certain flowers of Nymphaea subg. Hydrocallis. a. Termi
nal papilla cell of powdery stigma in N. rudgeana (Wiersema et al 2309). b. Subterminal papilla cell of
N. rudgeana (Wiersema et al. 2309). c. Spherical sclereid from anther of N. oxypetala (Fern?ndez
1299). Note embedded crystals of calcium oxalate. d. Acicular sclereid from anther of N. potamophila
(Madison et al. 6155). Bar = 10 um.
petals and stamens. Acicular sclereids may be found within the sepals, petals,
stamens, and carpellary appendages, or even projecting from the stigmatic ray
tissues in a few species. They are thick and needlelike in many species, but in
others they are thinner or more filiform. In the stamens of N. oxypetala unusual
spherical sclereids (Fig. lc) are apparently released (in dried anthers at least)
from the staminal interior along three sutures lateral to and between the anther
sacs. They were studied by Chifflot (1902) but their functional significance is
unknown.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 11
SEED MORPHOLOGY
According to Conard (1905), the seeds of water lilies display marked charac
teristics which can be of systematic value. They were the object of a study of the
Nymphaeaceae (sensu lato) by Weberbauer (1894), whose findings regarding
Nymphaea are discussed by Conard. The seed coat in Nymphaea has generally
been regarded as bitegmic (Chifflot 1902; Conard 1905; Khanna 1967). The outer
most cell layer is composed of lignified cells (sclereids), whose interdigitating
radial cell walls form a puzzlelike network over the surface of the seed. These
sclereids are, for the most part, arranged in longitudinal rows. They may be
almost isodiametric or squarish, but more frequently are elongated in a direction
perpendicular to the rows, except along the raphe where the elongation is in the
direction of the rows. The raphe may form a ridge of varying prominence along
one side of the seed. Other longitudinal ridges may be formed by the intercalating
ends of adjacent rows of sclereids being alternately raised up or level. The rows,
and the ridges when present, are interrupted as they converge toward both the
chalazal and micropylar ends of the seed. The outer tangential surface of the
sclereids may be variously supplied with papular outgrowths as well. These hair
like structures are, according to Conard, outgrowths of the radial cell walls of the
sclereids.
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12 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Seed Size and Color. Seeds of Nymphaea subg. Hydrocallis range from nearly
orbicular in N. jamesoniana and N. oxypetala to highly ellipsoid in some popula
tions of N. amazonum. With the exception of the relatively large seeds of N.
rudgeana, seeds of subg. Hydrocallis are smaller than those of other species of
Nymphaea. The smallest are those of N. jamesoniana, which barely exceed 0.5
mm in diameter. For some species, such as N. tenerinervia and N. glandulifera,
seed size and shape are reasonably uniform throughout the distributional range.
In others, such as N. conardii and especially N. amazonum and N. ampia of subg.
Brachyceras, both size and shape vary from one region to another. Seed material
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 13
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14 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 2. Light and scanning electron micrographs of seeds of Nymphaea subg. Hydrocallis. a.
SEM of whole seed, illustrating short, solitary hairs and highly intercalated, rectangular surface
sclereids with granulate fine surface topography, b. SEM of whole seed, illustrating medium-length,
clumped hairs with less intercalated, more isodiametric sclereids having a smooth topography, c. Near
median longitudinal section of seed of N. novogranatensis (Wiersema & Gonz?lez 2234) showing
overall seed anatomy; o = outer integument, i = inner integument, p = perisperm, a = aril, mr =
micropylar ridge, op = operculum, pi = plumule, c = cotyledon, e = endosperm. Bar = 50 |xm.
the presence of lignin. The outer integument is often detached from the inner
upon drying of the seed by breakdown of these thin-walled cells and the inner cell
walls of the sclereid coat. The inner integument, which stains a deep red with
safranin, appears as an amorphous band, although Chifflot (1902) indicated it was
composed of two layers of cells. Lying just within the inner integument, a narrow
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 15
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FIG. 3. Group I and II seeds of Nymphaea subg. Hydrocallis. a-c. Group I, N. rudgeana. d-f.
Group II, N. lasiophylla. g-i. Group II, N. lingulata. a, d, g. SEM of whole seeds, showing arrange
ment of papillae along alternate cell junctures (a. Wiersema et al. 2299. d. Wiersema et al. 2316. g.
Wiersema et al. 2308.). b, e, h. Light micrographs of seed coat, lacking granulate surface, with fewer
pit canals (arrow) in b and numerous pit canals in e and h. c, f, i. Light micrographs of seed coat
section. Note thin outer tangential wall and thick inner integument in c and thicker inner tangential
walls (arrow) of surface sclereids in f and i; o = outer integument, i = inner integument. Horizontal
bar = 0.1 mm, vertical bar = 20 \im.
16
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FIG. 4. Group Ilia seeds o? Nymphaea subg. Hydrocallis. a-c. N. amazonum subsp. amazonum.
d-f. N. tenerinervia. g-i. N. gardneriana. a, d, g. SEM of whole seeds, showing arrangement of
papillae along alternate cell junctures (a. Wiersema et al. 2304. d. Wiersema et al. 2307. g. Wiersema
2247a x 2214). b, e, h. Light micrographs of seed coat, displaying granulate surface, c, f, i. Light
micrographs of seed coat section showing details of testal anatomy; o = outer integument, i = inner
integument, p = perisperm, a = aril, t = transparent layer surrounding perisperm, arrow = pit canal.
Horizontal bar = 0.1 mm, vertical bar = 20 urn.
17
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18 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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FIG. 5. Group Illb and IIIc seeds of Nymphaea subg. Hydrocallis. a-c. Group Illb, N. novo
granatensis. d-f. Group Illb, N. conardii. g-i. Group IIIc, N. jamesoniana. a, d, g. SEM of whole
seeds (a. Wiersema & Gonz?lez 2234. d. Wiersema & Gonz?lez 2214. g. Wiersema et al. 2277). b, e, h.
Light micrographs of seed coat, displaying granulate surface, c, f, i. Light micrographs of seed coat
section. Note thinner outer tangenital wall of testa and surface papillae (arrow) in f. Horizontal bar =
0.1 mm, vertical bar = 20 ^m.
19
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t*>i ^ US'* JF
FIG. 6. Group IV seeds of Nymphaea subg. Hydrocallis. a-c. N. oxypetala. d-f. N. glandulifera.
g-i. N. potamophila. a, d, g. SEM of whole seeds, showing clumped distribution of hairs (a. Wier
sema & Gonz?lez 2232. d. Liesner 2892. g. Fr?es 22686). b, e, h. Light micrographs of seed coat, with
numerous pit canals (arrow) and lacking granulate surface, c, f, i. Light micrographs of seed coat
section, showing pit canals with dark inclusions (arrows). Horizontal bar = 0.1 mm, vertical bar = 20
urn.
20
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 21
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FIG. 8. Seeds of other subgenera of Nymphaea. a-c. N. odorata of subg. Nymphaea. d-f. N.
elegans of subg. Brachyceras, g-i. N. ampia of subg. Brachyceras, a, d, g. SEM of whole seeds (a.
Wiersema 342. d. Pringle 1955. g. Wiersema 2204a). b, e, h. Light micrographs of seed coat with
numerous pit canals and lacking granulate surface, c, f, i. Light micrographs of seed coat section. Note
thicker inner integument in i and additional cell layers present in outer integument of c. Horizontal
bar = 0.1 mm, vertical bar = 20 urn.
22
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 23
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24 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 9. Seed of Nymphaea lotus of Nymphaea subg. Lotos, a. SEM of whole seed (Wiersema
1992). b. Light micrograph of seed coat. Note virtual absence of pit canals, c. Light micrograph of
seed coat section. Note thickness of inner integument and relatively thin outer tangential wall of
surface sclereids. Horizontal bar = 0.1 mm, vertical bar = 20 urn.
arrangement of surface sclereids and granulate fine surface topography. This lat
ter feature, although it seems to be restricted within Nymphaea to subg. Hydro
callis, bears a strong resemblance to the cuticular papillae reported by Schneider
and Ford (1978) on seeds of the closely related genus Ondinea and to the granu
late surface observed by Collinson (1980) on Brasenia Schreber seeds. If one
assumes Conard's phylogenetic scheme for Nymphaea to be reasonably accurate,
a further examination of seeds of subgenera Brachy ceras, Lotos, and Anecphya
would be helpful in defining the primitive vs. derived status of this character
within Nymphaea.
The trend toward smaller seeds with shorter and more randomly distributed
hairs in Groups Illb and IIIc, however, would have to be considered advanced
within the subgenus. Other probable derived characters are the clumped distribu
tion of hairs and the more isodiametric surface sclereid shape evidenced in Group
IV seeds.
A final note should be made regarding the absence of two other species of
Nymphaea subg. Hydrocallis from this phase of investigation. These are N. belo
phylla and N. prolif?ra, both excluded due to a lack of available seed material at
the time of the initial study. In the former case this could be attributed to the
scarcity of collections of this species; however, in the latter case it was due to a
total preponderance of asexual reproduction, thereby circumventing normal seed
production, in all populations encountered thus far. On the basis of other mor
phological considerations, N. belophylla would be expected to display seeds akin
to those of Group IV. It is now possible to characterize seeds of N. prolif?ra,
however; those produced from a recent artificial hybridization display the charac
teristics of Group Ilia, as predicted by Wiersema (1984b).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 25
POLLEN MORPHOLOGY
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26 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
The pollen grains of subg. Hydrocallis possess the distinctive bandlike aper
ture characteristic of other Nymphaea pollen. According to Walker's (1974) classi
fication of zonasulculate pollen, they vary from anazonasulculate to zonizonasul
culate depending on the amount of expansion or compression of the operculum.
In subg. Hydrocallis the operculum bears a doughnut-shaped ring around its
perimeter. This ring seems to be produced from the constriction of the inner
portion of the operculum, with the dilated outer region delimited by a shallow
furrow. A puckered surface may develop over this constricted portion; otherwise,
the pollen grain is completely smooth. In overall shape it is reminiscent of a tiny
hamburger bun.
Only slight differences are detectable among the various species of subg.
Hydrocallis with regard to pollen morphology (Figs. lOa-i, lla-c). Differences
exist mainly in pollen size and in the extent of demarcation of the doughnut
shaped ring. This ring is fairly conspicuous in all species but is only slightly
evident in N. jamesoniana (Fig. lie). Pollen size is greatest in N. lingulata, N.
rudgeana, and N. lasiophylla, in decreasing order, with the other species observed
being fairly uniform in size. Confident determinations of most species of subg.
Hydrocallis are not possible, and little illumination of phylogenetic relationships
within the subgenus is provided from pollen analysis.
The greatest utility of pollen analysis has been in assessing the relationship of
subg. Hydrocallis to other subgenera of Nymphaea. As already indicated, the
pollen of subg. Nymphaea is highly ornamented and anazonasulculate. Pollen of
N. odorata (Fig. lid) and N. mexicana (Fig. lie) exemplify this morphology,
which appears very different from that of subg. Hydrocallis. In pollen of N.
mexicana, however, the operculum is patterned in a manner similar to that found
in subg. Hydrocallis. The inner portion is comparatively smooth, but along its
margin a dilated verrucate band is observed which may be homologous to the
doughnutlike ring evident in subg. Hydrocallis. This ring is clearly evident from
an examination of pollen of N. lotus of subg. Lotos (Fig. llf). Pollen of this
species also displays a smooth exine like that of subg. Hydrocallis. Pollen of subg.
Brachyceras, as evidenced in N. ampia (Fig. llg), N. elegans (Fig. llh), and N.
capensis Thunberg (Fig. Hi) shows no evidence of this ring whatsoever. The
relatively uniform pollen morphology exhibited by these three species and other
members of subg. Brachyceras that have been reported is characterized by a
rather broad equatorial furrow (zonizonasulculate), virtually identical hemispheric
portions, and a slightly roughened or puckered surface. Pollen of subg. Anecphya
was not available for examination but from published accounts (Walker & Doyle
1975) appears identical to pollen of subg. Brachyceras.
Pollen of subg. Lotos appears most similar to that of subg. Hydrocallis. The
close similarity observed between the two subgenera suggests a common origin for
this pollen type. Additional evidence for such a viewpoint will be discussed under
phylogeny.
CHROMOSOME NUMBERS
Published chromosome counts on other species of Nymphaea have indicated a
base number of x = 14 for the genus. The majority of counts were obtained by
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 27
FIG. 10. Scanning electron micrographs of pollen of nine species of Nymphaea subg. Hydrocallis.
a. N. novogranatensis (Wiersema & Gonz?lez 2234). b. N. glandulifera (Standley 73128). c. N. oxype
tala (Fern?ndez 1299). d. N. tenerinervia (Wiersema et al. 2307). e. N. lasiophylla (Wiersema et al.
2317). f. N. lingulata (Wiersema et al. 2308). g. N. rudgeana (Wiersema & Gonz?lez 2205). h. N.
amazonum subsp. amazonum (Wiersema 1400). i. N. prolif?ra (Asplund 16024). Bar = 5 \im.
Langlet and S?derberg (1927) and Gupta (1978, 1980); at the time of Gupta's
(1980) report counts were available for 22 of the ca. 40 species. Among the
apocarpous groups of Nymphaea, counts of 2n = 28, 56, and 84 have been
reported within subg. Brachyceras and 2n = 224 for the monotypic subg. Anec
phya. Within the syncarpous line counts of 2n = 56, 84, and 112 (and a number of
aneuploid counts: 64, 96, 105, 120, 160) have been reported for subg. Nymphaea,
and 2n = 28, 56, 84, and 112 for subg. Lotos. This evidence suggests that all other
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28 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 11. Scanning electron micrographs of pollen of subgenera Hydrocallis, Nymphaea, Lotos,
and Brachyceras of Nymphaea. a-c. Pollen of subg. Hydrocallis. a. N. conardii (Wiersema & Gonz?lez
2214). b. N. gardneriana (Wiersema & Gonz?lez 2210). c. N. jamesoniana (Wiersema 2031). d-e.
Pollen of subg. Nymphaea. d. N. odorata (Wiersema 784). e. N. mexicana (Wiersema 281). f. Pollen of
N. lotus of subg. Lotos (Wiersema 1442). g-i. Pollen of subg. Brachyceras. g. N. ampia (Wiersema &
Gonz?lez 2204a). h. N. elegans (Wiersema 1981). i. N. capensis (Robinson et al. 44). Bar = 5 um.
subgenera appear to share this base number. No species of subg. Hydrocallis had
been previously counted.
Successful mitotic counts were obtained from 24 populations, representing 11
of the 14 recognized species of subg. Hydrocallis and 2 probable hybrids. One
putative hybrid population (Wiersema et al. 2296) failed to yield an acceptable
count due to insufficient material. Those counts obtained are indicated in Appen
dix 1. Consistency of counts was observed for all populations within each species.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 29
On the basis of the observed counts some obvious relationships within the
subgenus become apparent. From information already discussed the base number
within Hydrocallis would be expected to be x = 14. Diploid counts of 2n = 28 for
N. jamesoniana, N. conardii, N. gardneriana, and N. novogranatensis, which are
documented in Figure 12a-d, support such an assumption. Seeming to contradict
this evidence is the existence of 2n = 18 in N. amazonum, N. prolif?ra, N.
lasiophylla, and N. lingulata (Fig. 12e-h).
The disparity in the two chromosome lines is believed to be the result of a
series of aneuploid alterations, thereby reducing the haploid number from 14 to 9.
Similar reductions in chromosome number have been observed in other genera of
both plants (Stebbins 1971) and animals (White 1973). Such alterations, which
have been termed "centric fusions" (White 1973) or "Robertsonian fusions"
(Jones 1976), can be achieved via unequal reciprocal translocations, resulting in
the formation of a larger metacentric chromosome from two smaller acrocentric
ones, with subsequent fixation of this translocation in the homozygous condition.
Five stepwise fusions of this type would be required to account for the aneuploid
shift observed between the two chromosome lines of subg. Hydrocallis. The dif
ference in relative size of the chromosomes between the two groups supports such
an interpretation. In the 2n = 28 line, all chromosomes are of similar size, as
apparent in Figure 12a-d. The 2n = 18 group, however, exhibits 10 large and 8
smaller chromosomes (e.g., Fig. 12g, h), the ratio expected from five haploid
fusions. The existence of an intermediate count of 2n = 20 for N. tenerinervia
(Fig. 12i), a species which demonstrates morphological and phytochemical ties to
both chromosome lines, also agrees with this interpretation. In N. tenerinervia 8
large and 12 smaller chromosomes are observed, in conformity with the expecta
tion that four fusions would have been required in this species.
An alternative explanation would reverse the proposed direction of chromo
some evolution, deriving the 2n = 28 line from the more primitive 2n = 18 line
through a series of aneuploid increases. This would suggest a primitive status for
subg. Hydrocallis within Nymphaea, as all other subgenera appear to be based on
what would then be a more advanced 2n = 28 number. Such an interpretation is
not supported by the preponderance of morphological and anatomical evidence in
the genus.
Two other species of subg. Hydrocallis yielded distinctive chromosome num
bers. Nymphaea rudgeana is believed to have 2n = 42. Although numerous
chromosome squashes have been prepared of this species, the observation of
actively mitotic cells has proven more difficult than in other species. As only a few
such cells in the proper configuration have been observed, some reservation ac
companies this reported count.
The suggestion put forth in the discussion of seed morphology that N. rudgeana
is somehow related to subg. Lotos is not contradicted by the 2n = 42 count.
Members of subg. Lotos have thus far yielded counts of 2n = 28, 56, 84, and 112. A
rare hybridization event involving an n = 28 Lotos individual and an n = 14
Hydrocallis parent would result in an individual with 42 chromosomes. Additional
karyological research, particularly with meiotic preparations, or enzyme electro
phoretic study is needed to further explore this possibility. Although numerous
hybrids, both natural and artificial, have been documented in Nymphaea, none are
believed to exist from crosses involving two subgenera (Conard 1905; Wood 1959).
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30 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 12. Chromosomes of nine species and one subspecies of Nymphaea subg. Hydrocallis. a-d.
2n = 28. Note similar sizes of chromosomes, a. N. jamesoniana (Wiersema etal. 2277). b. N. onardii
(Wiersema & Gonzalez 2214). c. N. gardneriana (Wiersema 1403). d. N. novogranatensis (Wiersema &
Gonz?lez 2234). e-h. 2n = 18. Note 10 larger and 8 smaller chromosomes, especially in g and h. e. N.
amazonum subsp. pedersenii (Wiersema et al. 2270). f. N. prolif?ra (Wiersema et al. 2273). g. N.
lasiophylla (Wiersema et al. 2293). h. N. lingulata (Wiersema et al. 2308). i. 2n = 20, N. tenerinervia
(Wiersema et al. 2307). j. 2n = 84, N. oxyp?tala (Wiersema & Gonz?lez 2232). Bar = 5 um.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 31
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32 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
of pollination syndrome. The flowers of these species open for two, three, three,
and four successive days, respectively.
Much less information exists on the floral biology of the night-flowering
subgenera Lotos and Hydrocallis. Other than brief observations on certain mem
bers of subg. Lotos (Conard 1905; Mauve 1967; Mitra & Subramanyam 1982;
Wiersema 1982) most of the data relate to subg. Hydrocallis. Field observations
of floral biology have been made on N. rudgeana, N. glandulifera, and N.
amazonum (Cramer et al. 1975; Prance & Anderson 1976; Prance 1980). All
observations indicate a cantharophilous pollination syndrome involving scarab
beetles of the genus Cyclocephala Latreille, including C. verticalis Burmeister,
C. castanea Olivier, and C. mollis Endr?di. The beetles have been reported to
feed on the highly developed carpellary appendages in subg. Hydrocallis (Prance
1980; Meeuse & Schneider 1980). Schneider (1982a) suggests an evolutionary
relationship between the beetle-pollination syndrome and the presence of these
enlarged clubshaped appendages. The lack of observation on the presence of
stigmatic fluid in subg. Hydrocallis has further led Meeuse and Schneider (1980)
to speculate on the possible absence of this fluid in these beetle-pollinated
flowers. Some question also remains whether beetles are indeed trapped inside
flowers during the day; some observers (Cramer et al. 1975) report they are and
others (Prance & Anderson 1976; Prance 1980) insist they can readily escape but
may passively remain.
Numerous crosses have been conducted within Nymphaea. These have been
made largely for horticultural purposes and have not involved members of subg.
Hydrocallis (Henkel et al. 1907; G. H. Pring 1934a, 1934b; G. W. Pring 1960).
Countless horticultural and several natural hybrids are known to exist within
Nymphaea (Wood 1959).
Floral Biology
Successful flowering in cultivation was achieved with eight species (N. amazo
num, N. conardii, N. gardneriana, N. jamesoniana, N. lingulata, N. novogranaten
sis, N. prolif?ra, N. tenerinervia); Walter Pagels (pers. comm.) supplied additional
data on a ninth, N. glandulifera. Flowers usually open for two consecutive nights
(sometimes three in N. jamesoniana). All exhibit protogamy except N. lingulata,
in which the anthers were observed to dehisce the first night. A pool of stigmatic
fluid is present in all first-day flowers, as shown in Figure 13a.
Differences in floral behavior among the taxa are apparent, particularly in
times of opening and closure of first-day flowers and closure of second-day
flowers. All second-day flowers observed commence opening at dusk, regardless
of species, except those of N. lingulata which open 1-2 hours later. On the basis
of similarities in floral responses, certain species groupings are apparent. One
includes N. glandulifera, N. conardii, N. gardneriana, and N. jamesoniana.
Flowers of this group open and close in synchrony. All these species, except N.
gardneriana, exhibit abundant autogamous seed production. Two other species,
N. novogranatensis and N. tenerinervia, are also similar in their opening and
closing times for flowers both nights. Autogamous seed production was observed
in some flowers of N. novogranatensis but in none of N. tenerinervia. Flowers of
N. lingulata, N. prolif?ra, and N. amazonum are not similar to other species in
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 33
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34 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 14. Flower opening times for first- and second-day flowers of nine species of Nymphaea
subg. Hydrocallis.
tions on N. rudgeana (Caspary 1878; Conard 1905) support its inclusion in this
latter group as well.
With regard to those species of subg. Hydrocallis observed, a floral clock
(Fig. 14) can be constructed similar to that produced by Conard (1905) for Nym
phaea. Although some degree of temporal separation is apparent between flowers
of the two groups just mentioned, only in N. amazonum is complete temporal
isolation approached due to the delayed pollen release in second-day flowers.
However, first-day flowers of N. amazonum could still be pollinated with pollen
of other species whose second-day flowers remain open until dawn. The potential
for hybridization between most species thus is not greatly affected by flowering
time.
Several items relevant to pollination are also apparent. One concerns differ
ences in floral odors between various species. These differences might relate to
the attraction of different species of pollinators, an interesting speculation put
forth by Meeuse and Schneider (1980) which demands further investigation.
Flowers of the pre-midnight flowering group generally produce a stronger and
more pleasant odor. Considerable odor is emitted from an entire population, as
the latter can readily be detected from a car passing by at night. Another differ
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 35
ence among species relates to the degree of opening of flowers. As a rule, first
day flowers open internally only enough to allow a small passage, ca. 1 cm or so
in diameter, to the fluid-filled stigma (Fig. 13b, d). In flowers of Nymphaea
conardii and N. gardneriana, however, no such passage was perceived. In second
day flowers the inner perianth and stamens of N. novogranatensis and N. teneri
nervia were frequently observed in a closed position at the time first-day flowers
were open. Presumably an insect visitor in these latter species would be required
to force its way into the floral interior in its foraging, thus ensuring a liberal
dusting with pollen. In both species the inner petals are greatly reduced and
would offer little resistance to such an entry (Fig. 13c).
Insect visitors to flowers of subg. Hydrocallis have been observed and cap
tured on only two occasions in the field. Cydocephala verticalis was collected
from the type locality of N. conardii in Venezuela and C. aff. p?trida Burmeister
(males were needed for positive identification according to B. C. Ratcliffe, pers.
comm.) from a population of N. lasiophylla in northeastern Brazil. This latter
Cydocephala, although requiring further identification, differs from those previ
ously reported as pollinators of subg. Hydrocallis. Further evidence of beetles
feeding on carpellary appendages has been detected in N. oxypetala flowers of
Venezuela, although no beetles were observed at that time.
Adaptations for autogamy are observable in certain species. In some, such as
N. conardii and N. jamesoniana, the stigmatic fluid persists in second-day flowers
and the stigma remains receptive to the pollen now released. Although the stigma
is likewise covered with fluid in second-day N. gardneriana flowers, it is no longer
receptive, as evidenced from the failure of all crosses attempted on that day. Two
other species appear to be approaching cleistogamy. In N. jamesoniana, flowers
frequently fail to open the first night and are therefore effectively homogamous.
Those of many populations, as in Ecuador and Florida, open only slightly, the
outer parts spreading only to a vertical position. Autogamous seed production is
abundant. In N. lingulata flowers are also homogamous. They, too, open only
slightly, as the thick, coriaceous sepals prevent the petals and stamens from
further reflexing (Fig. 13d). Autogamous seed production is also evident. No
seeds were produced from any emasculated flowers of subg. Hydrocallis in the
absence of cross-pollination, which indicates autogamy and not agamospermy is
responsible for seed production in unemasculated flowers under these conditions.
Crossing Studies
The pollen fertility of the eight species examined was presented in Wiersema
(1984b). With the exception of a Brazilian population which exhibited less than
5% viability, all other populations have pollen fertility exceeding 75%. This Bra
zilian population is clearly of hybrid origin, probably involving N. lasiophylla and
N. amazonum subsp. amazonum.
An assortment of crosses was conducted among the various species; see table
1. Unfortunately most crosses were conducted among members of the 2n = 28
chromosome line?N. conardii, N. gardneriana, N. jamesoniana, and N. novo
granatensis?as more flowers of this group were available in cultivation. All of
these species exhibited a moderate degree of interfertility. Crosses between this
group and the 2n = 18 line were very limited in number, but only N. novograna
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Table 1. Percentage seed set from artificial crosses among several species of Nymphaea subg. Hydrocal
Nymphaea.
Pollen parent
N. N. N. N. N. N. N
Seed parent amazonum conardii gar
Ia
N. amazonum
2 1 2 1
N. conardii 10-20 60 20-50 40
3 1 3
N. gardneriana 20-40 80 20-50
N. jamesoniana
N. lingulata
1 1
N. novogranatensis 30 <10 40-80
1
N. prolif?ra <5
N. rudgeana
1
N. tenerinervia 20
N. lotus
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 37
tensis, the most profusely flowering species in cultivation, has indicated any de
gree of interfertility with this other line. Interestingly, some degree of seed set
was observed in crosses involving N. novogranatensis and most other available
species of subg. Hydrocallis, which supports a central phylogenetic position for
this species within the subgenus. The lowest interfertilities recorded for N. novo
granatensis involve crosses with N. lingulata and N. jamesoniana, two species
which may occupy more peripheral positions in the phylogenetic tree. No seed set
was observed from any crosses of species of subg. Hydrocallis with those of other
subgenera. Most involved N. lotus of subg. Lotos, although none of these crosses
included N. rudgeana, which is presumed to bear some relation to this other
subgenus.
FLAVONOID CHEMISTRY
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38 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 39
VOUCHER COUNTRY
0.25
_J_
0.55
>
0.85 No.a TAXON OF ORIGIN
AMP LA Mexico
GARDNERIANA Argentina
JAMESONIANA Mexico
JAMESONIANA Ecuador
JAMESONIANA Florida, USA
JAMESONIANA Argentina
JAMESONIANA Brazil
GARDNERIANA Venezuela
GARDNERIANA Venezuela
GARDNERIANA Venezuela
GARDNERIANA Venezuela
MEXICANA Alabama, USA
AMP LA Florida, USA
AMP LA Ecuador
OXYPETALA Venezuela
ELEGANS Florida, USA
ODORATA Alabama, USA
CANDIDA West Germany
LINGULATA Brazil
LINGULATA Brazil
LOTUS Florida, USA
AMAZONUM Argentina
RUDGEANA Venezuela
RUDGEANA Brazil
RUDGEANA Brazil
AMAZ. X LAS10. Brazil
AMAZONUM Ecuador
TENERINERVIA Brazil
AMAZONUM Venezuela
AMAZONUM Ecuador
AMAZONUM Brazil
TENERINERVIA Brazil
TENERINERVIA Brazil
TENERINERVIA Brazil
PROLIFERA Ecuador
PROLIFERA Argentina
PROLIFERA Argentina
PROL. X AMAZ. ? Argentina
PROL. X AMAZ. ? Argentina
PROL. X AMAZ. ? Argentina
PROLIFERA Argentina
PROL. X AMAZ. ? Argentina
NOVOGRANATENSIS Venezuela
NOVOGRANATENSIS Venezuela
NOVOGRANATENSIS Venezuela
NOVOGRANATENSIS Venezuela
AMAZONUM Venezuela
LASIOPHYLLA Brazil
LASIOPHYLLA Brazil
CONARDII Venezuela
CONARDII Nicaragua
CONARDII Nicaragua
CONARDII Brazil
CONARDII Venezuela
Collection numbers are ofH.
J.Wiersema, except Nos. 8588 and 8612 which are
of R. R. Haynes.
FIG. 15. Phenogram derived from presence/absence data for 56 compounds isolated from
ions of Nymphaea.
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40 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
VOUCHER COUNTRY
0.25 0.55 0.85 No.* TAXON OF ORIGIN
1345 AMPIA Mexico
-2021 AMPLA Ecuador
1452 AMPLA Florida, USA
274 ODORATA Alabama, USA
1981 ELEGANS Florida, USA
?2196 CANDIDA West Germany
'1403 GARDNERIANA Venezuela
1428 GARDNERIANA Venezuela
2210 GARDNERIANA Venezuela
2247 GARDNERIANA Argentina
-1343 JAMESONIANA Mexico
-2030 JAMESONIANA Ecuador
?2032 JAMESONIANA Florida, USA
2265 JAMESONIANA Argentina
2290 JAMESONIANA Brazil
2232 OXYPETALA Venezuela
1412 GARDNERIANA Venezuela
281 MEXICANA Alabama, USA
i-]1411 NOVOGRANATENSIS Venezuela
2234 NOVOGRANATENSIS Venezuela
1420 NOVOGRANATENSIS Venezuela
1404 NOVOGRANATENSIS Venezuela
2028 PROLIFERA Ecuador
2241 PROLIFERA Argentina
2273 PROLIFERA Argentina
2238 PROL. X AMAZ. ? Argentina
2255 PROLIFERA Argentina
2254 PROL. X AMAZ. ? Argentina
2249 PROL. X AMAZ. ? Argentina
?2240 PROL. X AMAZ. ? Argentina
Ecuador
2013 AMAZONUM Ecuador
E2027 AMAZONUM
2291 AMAZONUM Brazil
Brazil
2314 TENERINERVIA Brazil
E2307 TENERINERVIA
2311 TENERINERVIA
? 2310 TENERINERVIA
Brazil
Brazil
-1436 AMAZONUM Venezuela
? 2296 AMAZ. X LASIO. Brazil
-2270 AMAZONUM Argentina
-2221 AMAZONUM Venezuela
? 2205 RUDGEANA Venezuela
2285 RUDGEANA Brazil
2299 RUDGEANA Brazil
1442 LOTUS Florida, USA
2214 CONARDII Venezuela
8588 CONARDII Nicaragua
8612 CONARDII Nicaragua
2230 CONARDII Venezuela
2289 CONARDII Brazil
2293 LASIOPHYLLA Brazil
2316 LASIOPHYLLA Brazil
2283 LINGULATA Brazil
2308 LINGULATA Brazil
8588 Nos.
Collection numbers are of J. H. Wiersema, except and 8612 which are
of R. R. Haynes.
FIG. 16. Phenogram derived from presence/absence data for 11 aglycones and 9 glyc
linkages of 54 populations of Nymphaea.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 41
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42 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 43
lated with aromatic acids are also present in the exceedingly complex flavonoid
profile of N. lasiophylla.
The similarity between N. lasiophylla and N. lingulata is apparent from an
analysis of their aglycones and glycosidic combinations. Both are characterized by
the predominance of 3-methylated forms of quercetin and myricetin. Between
the two species, only in N. lasiophylla have nonmethylated forms of these agly
cones been detected. As a result of this 3-methylation, glycosylation in both
species is restricted to 7- and 3'-glucosides or 3'-xylosides. Glycosylation at the
7 position is unique to these two species among the representatives of Nymphaea
examined.
Although 3-methylation is lacking among other species of subg. Hydrocallis,
it is evident in several other Nymphaea species, including N. elegans and N. ampia
of subg. Brachyceras, and N. candida and N. odorata of subg. Nymphaea. All
four species and N. lotus of subg. Lotos characteristically produce 3'-xylosides as
well. This latter feature is not restricted to N. lasiophylla and N. lingulata within
subg. Hydrocallis, since it is also present in N. tenerinervia and in some popula
tions of N. novogranatensis (in low levels) and N. amazonum. Congruity of sev
eral 3-methylated compounds underlies the clustering of N. lingulata and mem
bers of subg. Brachyceras and subg. Nymphaea in Figure 15. The association of
N. lingulata with these taxa suggests a relationship that will be elaborated on in
the discussion of phylogeny.
The flavonoids of two additional species of subg. Hydrocallis remain to be
discussed. One of these is N. rudgeana, whose flavonoid profile is dominated by
C-glycosylflavones and quercetrin glycosides, including the 3'- and 4'-glucoside.
Both of these latter compounds are present in certain populations of N. amazo
num as already mentioned; the 4'-glucoside was not observed elsewhere in the
genus. With the exception of quercetrin, the most concentrated flavonoid in N.
rudgeana, each compound detected in N. rudgeana is also present in the Argen
tine N. amazonum subsp. pedersenii, indicating a possible affinity between these
two taxa. Flavonoid evidence does not, however, clarify the relationship between
N. rudgeana and N. lotus of subg. Lotos that was postulated from morphological
and seed evidence. Both similarities and differences exist in the flavonoid profiles
of the two species. Further phytochemical analysis of other members of subg.
Lotos could aid in resolving this dilemma.
The remaining species of subg. Hydrocallis examined, N. oxypetala, has a
flavonoid profile which combines elements of some of the other groups. The
absence of C-glycosylflavones and the presence of quercetrin align N. oxypetala
with species of the first group. However, the presence of Compounds A, B, and C
in N. oxypetala together with myricetin derivatives suggests a linkage with the
second group. Unfortunately three additional species of subg. Hydrocallis, N.
glandulifera, N. belophylla, and N. potamophila, which on morphological grounds
appear to be most closely allied to N. oxypetala, were not available for chemical
examination.
Hybridization. A number of putative hybrid populations were also examined.
All of these populations reproduced exclusively by asexual means and were as
sumed to be incapable of sexual reproduction. Chromosome counts obtained from
two such populations indicated them to be triploid. Both of these and three
additional populations, all from Argentina, cluster with N. prolif?ra, one of the
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44 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 45
PHYLOGENY
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Table 2. Character states of subgenera of Nymph
Position of anthers
on outer stamens unknown marginal medial
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 47
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48 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
sclereids, and polyploid nature, is perhaps the most specialized of all members of
subg. Hydrocallis. The position of N. belophylla remains uncertain at this time;
the possibility that N. belophylla represents a recent hybrid between two of these
species cannot be eliminated.
Although N. novogranatensis appears to approximate the ancestral form
within the 2n = 28 line, it also most closely resembles the 2n = 18 group. Its
closest relative is N. tenerinervia, both in morphology and floral biology. Nym
phaea tenerinervia, 2n = 20, is closely allied phytochemically to N. amazonum of
the 2n = 18 line and may represent a link between the two major groups. All
three of these species exhibit most of the primitive features discussed above and
are believed to be associated with the emergence of this additional chromosome
line. The other three members of the 2n = 18 line, N. prolif?ra, N. lasiophylla,
and N. lingulata, are similar in lacking the tetrameric arrangement of flower parts
present in other species of subg. Hydrocallis. Nymphaea prolif?ra and N. lasio
phylla are further related as the only members of subg. Hydrocallis which produce
tuberiferous flowers, but this feature is absent from N. lingulata. Among these
three species, N. prolif?ra is most similar to N. amazonum with regard to overall
morphology and phytochemistry. Nymphaea lasiophylla and N. lingulata, the only
members of subg. Hydrocallis with tapering carpellary appendages, are obviously
closely related, their relationship further substantiated by distinctive seed mor
phologies and flavonoid profiles.
Several features which might be presumed to be ancestral due to their pres
ence in other subgenera are absent from most species of subg. Hydrocallis but
present in N. lingulata and N. lasiophylla. In particular, the less ordered arrange
ment of the perianth, absence of petaloid stamens, tapered carpellary append
ages, certain seed characteristics, and presence of 3-methylated flavonoids in
these two species fit this pattern. However, the probable pathway for the emer
gence of the 2n = 18 chromosome line, as already discussed, would seem to have
involved an ancestor similar to N. novogranatensis, N. tenerinervia, and N.
amazonum, which do not display these characteristics. This supposed pathway
would dictate that N. lingulata and N. lasiophylla be viewed as more advanced
within the 2n = 18 line. Assuming this phylogenetic interpretation to be correct,
the features in question must have been acquired secondarily by N. lingulata and
N. lasiophylla. The possibility that this acquisition may be the result of genetic
interaction with other subgenera remains to be investigated.
Intersubgeneric hybridization with subg. Lotos may have been involved in the
formation of N. rudgeana; however, enough evidence has not yet been gathered
to confirm this hypothesis. The relationship of N. rudgeana to subg. Lotos has
been discussed in several sections above. Within subg. Hydrocallis, N. rudgeana is
most closely allied phytochemically to N. amazonum subsp. pedersenii.
TAXONOMY
Nymphaea Linnaeus, Sp. pi. 510. 1753, nom. cons. Leuconymphaea Ludwig ex
Kuntze, Revis, gen. pi. 1: 11. 1891.?Type: Nymphaea alba Linnaeus
(typ. cons.).
Castalia Salisbury, Ann. Bot. (K?nig & Sims) 2: 71. 1806.?Lectotype, here
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 49
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50 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
1. Carpellary appendages Ungulate-tapering; transition from petals to stamens abrupt, the outer
stamens only slightly or not at all petaloid.
2. Petals 20-26; tuberiferous flowers usually present; acicular sclereids absent from leaf
mesophyll. 12. N. lasiophylla.
2. Petals 8-14; tuberiferous flowers absent; acicular sclereids common in leaf mesophyll.
13. N. lingulata.
1. Carpellary appendages linear to strongly cl?vate; transition from petals to stamens gradual,
the outer stamens distinctly petaloid.
3. Leaf blades irregularly dentate; seeds 1.2-2.4 mm long. 14. N. rudgeana.
3. Leaf blades entire; seeds rarely exceeding 1.3 mm long.
4. Mature leaves mainly submersed (if floating then reduced in size), lacking sclereids in
mesophyll; sepals and outer petals acuminate. 7. N. oxypetala.
4. Mature leaves floating, sclereids common in mesophyll; sepals and outer petals
rounded to acute.
5. Leaf blades sagittate, over 1.8 times as long as wide.
6. Carpellary appendages 10-17 mm long; acicular sclereids present in leaf meso
phyll and in stigmatic rays. 5. N. belophylla.
6. Carpellary appendages ca. 7-9 mm long; acicular sclereids absent from leaf
mesophyll and from stigmatic rays. 6. N. potamophila.
5. Leaf blades suborbicular to elliptic, less than 1.6 times as long as wide.
7. Leaf veins with slightly to strongly evident cross veins centrally, forming weblike
venation pattern; stigmatic rays often containing few to numerous acicular scle
reids.
8. Carpellary appendages 3-9 mm long, less than Vs times length of longest
sepal; seeds 1.1-1.2 times as long as broad. 1. N. jamesoniana.
8. Carpellary appendages mostly 9 mm or more long, the longest more than Vs
times length of longest sepal; seeds 1.2 or more times as long as broad.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 51
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52 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
sclereids; undersurface green, sometimes with darker flecks, when dried usually
marked with slender, branched, rusty brown lines; venation conspicuously web
like centrally, dichotomous and faintly reticulate toward margin, the principal
veins 11-17. Petioles to 5 mm in diameter, glabrous, with 2 major and 4-6 minor
air canals. Peduncles to 7.5 mm in diameter, glabrous, with an inner ring of 6-7
major air canals surrounded by 12-14 minor ones. Flowers floating, the sepals,
petals, and outer stamens in distinct whorls of four. Sepals 2.5-6.5 cm long, 1.1
3.5 cm wide, uniformly green, ovate, acute to somewhat rounded at apex, when
dry marked with slender rusty brown lines, containing numerous acicular scle
reids, inconspicuously nerved. Petals 12, 16, or 20, containing acicular sclereids,
the outer greenish at least abaxially, mostly equal to or somewhat shorter and
narrower than sepals, with apex acute to rounded, the inner 1.8-4.5 cm long,
0.7-2 cm wide, creamy white, acute or rarely obtuse; transition to stamens
gradual. Stamens 35-85, creamy white, containing acicular sclereids, the outer
most to 3.9 cm long, with filaments to 15 mm wide, the inner with filaments to 2.5
mm wide and connectives prolonged to 0.3 (-0.5) mm beyond anthers, the inner
anthers 0.36-0.65 times as long as stamen; pollen diameter 26-32 |im. Carpels
19-33, the appendages 3-9 mm long, cream-colored with somewhat reddish or
purplish tips, the tips slightly cl?vate, 0.6-1.5 mm in diameter, the stigmatic rays
with none or few, occasionally numerous, projecting acicular sclereids and papil
lae cells remaining attached; axial process mostly 2-3 mm long, broadly acute to
acuminate. Fruits usually present, to 8 cm in diameter; seeds 0.5-0.8 mm long,
0.45-0.7 mm in diameter, 1.1-1.2 times as long as broad, brownish or whitish
when dry, with numerous short hairs 10-50 \im long, the surface finely granulate.
Tuberiferous flowers absent. Chromosome number: 2n = 28. Figs. 5, 11, 12, 17.
Phenology. Collected in flower and fruit throughout the year in tropical areas
and during warmer seasons in subtropical areas.
Distribution. (Fig. 18). Tropical and subtropical America, including Florida,
Mexico, Central America, Colombia, Ecuador, Brazil, Paraguay, and Argentina;
in freshwater ditches, ponds, or slow-moving streams, often with Characeae.
Specimens Examined. U.S.A. Florida: Citrus Co., ford leading to Sheep Island, "The Cove,"
Floral City, 1 Oct 1940, St. John s.n. (FLAS); DeSoto Co., 2.5 mi NW of Arcadia, 4 Nov 1967,
Conard s.n. (FLAS, USF); Lee Co., ca. 7 mi E Ft. Myers, Les 114 (USF), Les 140 (USF); Ft. Myers,
Les 159 (USF); Levy Co., Lebanon Station, Cooley et al. 7151 (FLAS, USF); Sarasota Co., Fla 72,
0.4 mi E Myakka R., Wiersema 2032 (UNA, US, USF).?Mexico. Chiapas: Mpio Arriaga, 6 km N
Arriaga, Breedlove 26918 (ENCB); Mpio Acala, road to La Angostura, 12-14 km S Tuxtla, Breedlove
37370 (MO), Mpio Tonal?, NW Puerto Arista, Breedlove & Thome 20917 (ENCB, MO, NY). Ta
basco: N-20, 4 km E Poblado el Naranjito, Men?ndez 255 (BM, K, MO). Tamaulipas: Mex-180, 14
km N of R?o Carrizal, Haynes 5441 (UNA), Wiersema 1341 (F, UNA, US). Veracruz: Mpio Paso de
Ovejas, Mata Mateo, Ventura 7681 (ENCB).
El Salvador. La Uni?n: Lago del Olomega, 8 km W La Uni?n, Fassett 28615 (F, WIS). San
Miguel: N end of Laguna Jocotal, San Salvador, Fassett 28565 (F, WIS).?Honduras. Yoro: 3 km N
of Las Lomas, Lentz 1187 (EAP, UNA).?Nicaragua. Matagalpa: 8 km E San Isidro, Haynes 8572
(MO, UNA). Rivas: Las Piedras, Haynes 8430 (MO, UNA). Prov. unkown: vicinity of Managua, lago
de Granada, Gamier 298 (US).
Argentina. Chaco: Depto La Leonesa, Ruta 11 between Resistencia and Formosa, Wiersema &
Vanni 2265 (CTES, UNA, CORD). Corrientes: Depto Mburucuy?, Estancia "Santa Teresa," Peder
sen 7741 (C); Depto San Miguel, Ruta 5, 9 km SE Ca? Cat?, Wiersema et al. 2277 (CTES, UNA).
Formosa: Depto Piran?, 6 km NW Piran?, Morel 531 (LIL); Depto Patino, riacho Montelindo, Arenas
2060 (BACP, UNA). Salta: Depto Oran, Aguaray, Schreiter 11035 (LIL). Santa Fe: Depto Capital,
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 53
Laguna Setubal, Rinc?n Norte, Mart?nez-Achenbach 97, 159 (LIL). Tucum?n: Depto Mon
Acharal, Venturi 1701 (BAB, GH, LIL, US).?Brazil. Alagoas: BR-316 at km 172, 8 km E P
dos Indios, Wiersema et al. 2290 (INPA, IPA, MO, NY, UNA). Bahia: Rio Salitre, between J
and Sobradinho, Wiersema et al. 2300 (IPA, UNA). Cear?: BR-116 at km 65, 65 km S Fo
Wiersema et al. 2315 (IPA, UNA). Maranh?o: BR-316 at km 598, 23 km W Rio Parnaiba, W
et al. 2306 (IPA, UNA). Mato Grosso: Mun. Pocon?, Rodavia Transpantaneira, da Silv
(UFMT). Mato Grosso do Sul: Estrada Transpantaneira, Acampamento do Exercito, Heringe
868 (UB). Pernambuco: Mun. Boa Vista, BR-428 at km 76, Riacho das Gar?as, Wiersema et
(IPA, UNA). State unknown: S?o Marcos, L?tzelburg 20502 (M).?Colombia. Atl?ntico: re
Barranquilla, Elias 1387 (F, US).?Ecuador. El Oro: between M?chala and Santa Rosa, 6 Ju
Guldmann s.n. (K). Guayas: hwy 25 Guayaquil-Babahoyo, 17 km S Babahoyo, Wiersema et
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54 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
90?W 60?W
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 55
nights and dry the third night. Stamens of three-day flowers are exposed and
begin dehiscing the second night. Two-day flowers fail to open the first night of
this three-day sequence. In the Argentine plants the flowers open wide, the sepals
and petals becoming broadly reflexed; however, in plants cultivated from Ecuador
(Wiersema et al. 2026) and from Florida (Wiersema 2031) the flowers barely open,
the sepals and petals spreading only to a vertical position. All flowers emit a
strong aromatic fragrance. Crossing studies indicate cross-pollination is possible
both the first and second night for three-day flowers. Self-pollination is clearly
possible on the second day.
Reproduction in N. jamesoniana is entirely dependent on seed production.
No form of vegetative propagation was observed in field or cultivated plants. Seed
production appears to be largely autogamous, with fruits and seeds always devel
oping, even in the absence of potential pollinators.
Nymphaea jamesoniana has the smallest seeds with the shortest surface papil
lae in subg. Hydrocallis. Based on considerations of chromosome number, phyto
chemistry, overall morphology, and seed morphology, N. jamesoniana displays its
strongest affinity to N. conardii. Its shorter carpellary appendages readily sepa
rate it from this species.
Unusually robust forms have been observed both in Florida and Argentina.
Plants in Argentina and Paraguay have slightly shorter carpellary appendages and
more coriaceous leaves than those in other areas and have been treated as N.
gibertii. Seed morphology and phytochemistry of these plants is identical to that
observed in typical N. jamesoniana. Plants in northeastern Brazil produce larger
seeds than other N. jamesoniana populations and have slightly longer and more
cl?vate capellary appendages. Seeds of Mexican plants were also unusual in their
shorter, almost nonexistent, papillae.
The specimen from the Delessert Herbarium at G was chosen as the lectotype
of N. jamesoniana, as this specimen was cited in Planchon's expanded 1853 treat
ment along with material of the Herbarium Webbianum which I did not examine.
In describing his N. sagittariaefolia, Lehmann (1853) cited only: "in America
centrali collegit filius Guilielmus Lehmann." According to Caspary (1878) no such
specimen existed in Lehmann's herbarium, only the collection of Jameson which
was not mentioned in Lehmann's manuscript. Lehmann (1854) later indicated his
species to be synonymous with N. jamesoniana Planch?n; the collections of Jame
son from Lehmann's herbarium at B and at K bear the name N. sagittariaefolia in
Lehmann's handwriting.
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56 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
when dried with numerous tiny prominences from underlying trichosclereids inter
spersed with short elevated lines often visible from underlying acicular sclereids;
undersurface green, when dried often marked with slender, branched rusty brown
lines; venation slightly to evidently weblike centrally, dichotomous and faintly
reticulate toward margin, the principal veins 9-15. Petioles to 4 mm in diameter,
glabrous, with 2-4 major air canals and 4-6 minor ones. Peduncles to 6.5 mm in
diameter, glabrous, with an inner ring of 6 major air canals surrounded by 12
minor ones. Flowers floating, with sepals and petals in distinct whorls of four.
Sepals 3-6 (-7.2) cm long, 1-3 cm wide, uniformly green, oblong-ovate, acute or
somewhat rounded at apex, often marked with slender rusty brown lines when
dry, containing acicular sclereids, inconspicuously nerved. Petals usually 12, 16,
or 20, containing numerous acicular sclereids, the outer greenish abaxially,
shorter than sepals, with apex acute to somewhat rounded, the inner 2.1-4.4 cm
long, 0.6-2 cm wide, creamy white, acute to acuminate; transition to stamens
gradual. Stamens 24-80, creamy white, containing acicular sclereids, the outer
most to 3.3 cm long, with filaments to 15 mm wide, the inner with filaments to 1.5
mm wide and connectives prolonged to 0.3 mm beyond anthers, the inner anthers
0.4-0.7 times as long as stamen; pollen diameter 25-40 \im. Carpels 16-30, the
appendages 7-16 mm long, cream-colored, with cl?vate tips to 1.2 (-1.5) mm in
diameter which usually darken upon drying, the stigmatic rays with projecting
acicular sclereids and papillae cells remaining attached; axial process 1-1.5 mm
long and blunt. Maturing fruits usually present; seeds 0.65-0.9 mm long, 0.5-0.7
mm in diameter, 1.2-1.4 times as long as broad, with longitudinal rows of short
hairs 25-70 \im long, the surface finely granulate. Tuberiferous flowers absent.
Chromosome number: 2n = 28. Figs. 5, 11, 12, 19.
Phenology. Collected in flower and fruit from June (Brazil) or July (else
where) to December.
Distribution. (Fig. 20). Southern Mexico through Central America to north
ern South America, including Colombia, Venezuela, and northern Brazil; also
found in Cuba, Puerto Rico, and Hispaniola; mostly in temporary ditches or
ponds in inland habitats.
Representative Specimens. Cuba. Santa Clara: Yag?ey Chico, in swampy forest at Laguna del
Yanco, Ekman 17007 (G, S).?Dominican Republic Duarte: Pimental, Ekman 13277 (S).?Haiti.
Dept. du Nord: Bayeux, between Bord de la Mer and Morne ? Madame, Ekman 2679 (S).?Puerto
Rico. A?asco, near "Hatillo," Sintenis 5688 (BM, G, GH, GOET, K, M, P, S).
Mexico. Chiapas: Mpio Jiguipilas, 16 km ESE of Tierra and Libertad along road to Villa Flores,
Breedlove 27347 (MO).
Belize. Punta Gorda, Schipp 1026 (BM, G, F, K, MO, NY, S).?El Salvador. Santa Ana: 13
km N of Texistepeque, 28 km N of Santa Ana, Fassett 28279 (F, WIS).?Honduras. Comayagua: 5
km W of Comayagua, Molina & Molina 22704 (BM, F, G, S).?Nicaragua. Chontales: 2 Vi km SW of
Quebrada Niscala along road Acoyapa-San Carlos, Haynes 8588 (UNA). Le?n: Las Penitas, Haynes
8612 (UNA). Managua: Sierra de Managua, 1930-1940, Gamier s.n. (F). Rio San Juan: 0.5 km S R?o
Oyate along road to San Miguelito, Nee & Miller 27620 (MO, UNA).?Panama. Canal Zone: Barro
Colorado Island, Ebinger 553 (F). Dari?n: pasture belonging to Pablo Oth?n, Duke & Bristan 283
(MO). Panam?: between Panam? and Chepo, Dodge et al. 16712 (G, MO, P). San Blas: Ailigand?
area, Hammel & D'Arcy 5007 (MO).
Brazil. Alagoas: Mun. Marimbondo, km 218 of BR-316 in Rio Tapuia, Wiersema et al. 2289
(IPA, UNA). Para: Bel?m-Brasilia, Rio Marsh, Brantjes 707801 (SP).?Colombia. Guajira: R?oha
cha, Haught 4431 (COL, US). Santander: Puerto Wilches and vicinity, Killip & Smith 14821 (F, GH,
NY, US).?Venezuela. Apure: hwy 19 at km 44, W of Guasdualito, Wiersema & Gonz?lez 2230
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 57
FIG. 19. Nymphaea conardii (Wiersema & Gonz?lez 2214). a. Undersurface of leaf. b. Opened
flower, c. Unopened flower bud. d. Cross section of petiole, d. Cross section of peduncle.
(MY, UNA, US, VEN). Bol?var: on road to San Felix, between Upata and Altagracia, 1-10 km NW
Upata, Steyermark 57693 (F, VEN). Cojedes: Sistema de Riego Las Majaguas, Presas 1-2-3, Sector
Agua Blanca, Trujillo 11181 (MY). Portuguesa: Laguna El Sam?n, near Acarigua, Solazar 27 (US,
VEN). Tachira: between Piscuri and Ayari, Steyermark & Rabe 96622 (NY, US, VEN). Zulia:
carretera Machiques-Col?n, 15 km S Maracaibo, Steyermark & Fern?ndez 99560 (VEN).
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58 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
3. Nymphaea gardneriana Planch?n, FI. Serres Jard. Eur. 8: 120. 1852. Nymphaea
passiflora Lehmann, Hamburger Garten- Blumenzeitung 9: 205, 217.
1853, nom. superfl. Leuconymphaea gardneriana (Planch?n) Kuntze, Re
vis, gen. pi. 1: 11. 1891.?Type: "Brasilia, in prov. Piauhy," Jul-Sep
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 59
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60 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 21. Nymphaea gardneriana (Wiersema & Gonz?lez 2215). a. Undersurface of leaf. b.
Opened flower, c. Unopened flower bud. d. Cross section of petiole, e. Cross section of peduncle, f.
Upper surface of leaf.
Phenology. Collected in flower throughout the year in tropical areas, but only
during warmer seasons in subtropical regions further south; collections in fruit,
although quite rare, suggest a similar schedule for fruiting.
Distribution. (Fig. 18). South America, including Venezuela, Brazil, Bolivia,
Paraguay, and Argentina.
Specimens Examined. Argentina. Chaco: Margarita Bel?n, Aguilar 751, 1045 (LIL); Colonia
Ben?tez, Wiersema et al. 2247a (UNA). Corrientes: Depto Concepci?n, 28 km SE Concepci?n, Schi
nini et al. 13270 (CTES); Depto Itat?, Ruta 12, 60 km E Itat?, Ahumada et al. 876 (CTES). Formosa:
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 61
near Guayerles, Jorgensen 3124 (GH, LIL, MO, US); Depto Matacos, 1 km N Tolder?a Toba, Arenas
2272 (BACP, UNA). Misiones: Depto Capital, Posadas, Rodrigues 98 (BAB).?Bolivia. Santa Cruz:
Prov. Nuflo de Ch?vez, 20 km S Concepci?n, Krapovickas & Schinini 32094 (CTES); Prov. Chiquitos,
28 km NE San Jos?, Krapovickas & Schinini 36605 (CTES).?Brazil. Acre: Estrada Alemanha,
Cruzeiro do Sul, Prance et al. 11912 (NY, UNA). Goi?s: Mun. Yatai, Macedo 3320 (US); Mun.
Alvorada, BR-020 at Rio Macacos, Hatschbach 39372 (MBM). Mato Grosso: inter Guia et Cuiab?,
14 May 1894, Malme s.n. (S). Minas Gerais: S?o Jo?o del Rey, Schwacke 10015 (P). Para: Taperinha
bei Santar?m, Ginzberger 657 (F, W); locality unknown, 1832, Siber s.n. (BR). S?o Paulo: Moji
Gua?u, Mos?n 1102 (S).?Paraguay. Paraguar?: Plaine de Mbatobi, Balansa 521 (BM, BR, G,
GOET, K, S). San Pedro: Primavera, R?o Tapiracuai, Woolston 1257 (K, NY, US).?Venezuela.
Apure: Mantecal, Ramia 4540 (VEN), Ramia & Montes 4701 (VEN); S of Bruzual, 3 km N of Ca?o
Guaritico, Wiersema & Gonz?lez 2215 (COL, F, MY, UNA, VEN). Barinas: km 30 of hwy Barinas
Cuidad Bolivia, Trujillo 13407 (MY). Bol?var: Sabana Cardona, R?o alto Suapure, Puig 3050 (VEN).
Cojedes: 10 km S of San Carlos, Trujillo 13825 (MY). Gu?rico: estaci?n experimental "Los Llanos,"
Trujillo 8881 (MY); hwy 2, 2 km S R?o Gu?rico near Calabozo, Wiersema 1403 (F, MO, NY, UNA,
US, VEN); hwy 2, 53 km S Calabozo, Wiersema 1412 (UNA, VEN); hwy 2, 9.5 km S R?o Gu?rico
near Calabozo, Wiersema 1428 (UNA, VEN). Portuguesa: hwy 5, 17 km SW of Guanare, 2 km NE of
R?o Tucupido, Wiersema & Gonz?lez 2210 (MY, NY, UNA, VEN); Depto Guanare, terrenos de la
Unellez, Stergios 7077 (PORT, UNA).
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62 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
carpellary appendages, with the tips of these appendages more incurved than in
populations further north. Several Brazilian populations, as well as plants from
the type locality, exhibit leaves which are noticeably more peltate than those in
other parts of the range. There is evidence from some populations that this
characteristic may be variable, but unfortunately none of these populations has
been studied in the field or in cultivation by me. Overall, they display a similarity
to other populations in Argentina and Venezuela that have been scrutinized more
extensively. The results of this study, particularly with regard to seed morphology
and phytochemistry, support the acceptance of a single variable species.
The specimen at G from the herbarium Delessert has been chosen as the
lectotype of N. gardneriana; it was one of two specimens cited by Planch?n in his
expanded 1853 treatment. The name N. passiftora Lehmann is superfluous, be
cause Lehmann cited the type of N. gardneriana. The collection of C. Beyrich
from Brazil (Serra d'Estrella) also cited by Lehmann (1853) could not be located
by Caspary (1878) in any herbarium, including Lehmann's. Caspary questioned
the authenticity of this reported collection.
4. Nymphaea glandulifera Rodschied, Med. Bern. Kol. Rio Essequebo 76. 1794.
Nymphaea blanda G. Meyer, Prim. fl. esseq. 201. 1818, nom. superfl.
Castalia blanda (G. Meyer) Lawson, Proc. and Trans. Roy. Soc. Canada
6: 117. 1888. Leuconymphaea blanda (G. Meyer) Kuntze, Revis, gen. pi.
1: 11. 1891.?Type: Guyana. Rio Essequibo, Rodschied 155 (lectotype,
here designated: GOET!). The specimen labelled Rodschied 155 at
GOET contains one leaf of N. glandulifera. The flower, described by
both Rodschied and Meyer, which belongs with this leaf and which (fide
Caspary 1878) was once included with this leaf is mounted with Rodschied
281, the type of N. rudgeana G. Meyer.
N. fenzliana Lehmann, Hamburger Garten- Blumenzeitung 9: 205, 217. 1853.
N. blanda var. fenzliana (Lehmann) Caspary in Martius, Fl. bras. 4(2):
173. 1878.?Type: Guatemala. San Juan de Nicaragua, 1841, Friedrichs
thal 1878 (holotype: W, destroyed; isotype: K!). Although Lehmann cited
only "San Jouan de Nicaragua"?apparently taken from a specimen sent
to him by E. Fenzl of the University of Vienna?Caspary (1878) indicates
the cited collection to be that of Friedrichsthal at W and K.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 63
1.1-2.1 (-2.8) cm wide, green, oblong-ovate, acute at apex, when dry often
marked with slender rusty brown lines at base, containing acicular sclereids, in
conspicuously nerved. Petals 12-20, creamy white, containing acicular sclereids,
the outer slightly shorter than sepals, with apex acute, the inner 2-3.5 (-5) cm
long, 0.3-1.6 cm wide, acute to acuminate; transition to stamens gradual. Sta
mens 45-98, creamy white, containing acicular sclereids, the outermost to 3.7 cm
long, with filaments to 8 mm wide, the inner with filaments to 1.1 mm wide and
connectives prolonged to 0.4 mm beyond anthers, the inner anthers (0.4-) 0.5
0.7 times as long as stamen; pollen diameter ca. 36 \im. Carpels 19-39, the
appendages 7-14 mm long, cream-colored, linear to slightly cl?vate, with tips
usually darkening upon drying, to 1.2 mm in diameter, the stigmatic rays lacking
acicular sclereids and with papillae cells mostly remaining attached; axial process
blunt. Fruits normally present; seeds 0.8-1 mm long, 0.5-0.7 mm in diameter,
1.4-1.6 times as long as broad, dark green to dark reddish-brown when dry, with
clusters of long hairs 45-180 (xm long, the fine surface topography smooth. Tuber
iferous flowers absent. Chromosome number unknown. Figs. 6, 10, 22.
Phenology. Collected in flower and fruit throughout the year.
Distribution (Fig. 20). Northern South America and Central America, from
Amazonas, Brazil, west to northern Peru and Ecuador and north to Guatemala,
Belize, Trinidad, and French Guiana; not known from Panama or El Salvador.
Collections from Florida referred to this species (as N. blanda) by Ward (1977)
are N. jamesoniana.
Specimens Examined. Belize. Dist. Toledo, Crique Sarco, Hedger 326 (BM).?Costa Rica.
Prov. unkown: Marais de Cucurba, Estrella Calamanea, Fittier 9376 (BR, US); Monte Verde, Stork
1897 (F). Lim?n: Cahuita, Baker & Burger 26 (F, U). Puntarenas: Playa Blanca, Golfo Dulce, Valerio
408 (F); near Jalaca Station, between head of Golfo Dulce and Palmar Sur, Allen 6278 (F); Osa
Peninsula, Corcovado Nat. Park, Liesner 2892 (MO).?Guatemala. Izabal: near Puerto Barrios,
Standley 72857 (F), Standley 73128 (F); between Puerto Barrios and San Tom?s, 1 xh mi SE Puerto
Barrios, Steyermark 39867 (F).?Honduras. Atl?ntida: vicinity of Tela, Standley 55153 (F, US)?
Nicaragua. Zelaya: 2 km SW of Bluefields, Moreno 24967 (MO, UNA).
Brazil. Acre: Vicinity of Turauac?, France et al. 7350 (INPA, NY, UNA). Amazonas: km 5
Manaus-Porto Velho road, Berg et al. F17628 (INPA, NY, UNA); Jacaretinga, S of Rio Amazonas,
opposite Careiro Lake, France et al. 23302 (NY, UNA). Para: Cacaual Grande, Aguas Pr?tas, Black
52-15458 (IAN).?Colombia. Choc?: Trocha Coredo, Fuchs et al. 22176 (COL).?Ecuador. Los
R?os: km 56 of road Quevedo-Sto. Domingo, R?o Palenque Biol. Sta., Dodson 6172 (MO). Morona
Santiago: Lago de Kumpaka, 12 Aug 1982, Steinitz-Kannan s.n. (UNA), Steinitz-Kannan 409 (UNA).
Prov. unknown: Coast plain, Rimbach 86 (F), Rimbach 288 (US).?French Guiana. Near Kourou,
Schnell 11221 (P).?Guyana. Near Bartices, Essequibo River, Jenman 5746 (K, NY); locality un
known, Jenman 7773 (GH, K, NY), 1824, Parker s.n. (K).?Peru. Amazonas: Prov. Bagua, 208 km
by road NE of Chiriaco, Barbour 4408 (MO). Loreto: Iquitos, Killip & Smith 27173 (F, NY, US),
Asplund 14388 (G, S, US), Martin et al. 1614 (INPA).?Surinam. Brokopondo: Brownsweg, Cramer
LBB14915 (U). Marowijne: E of Perika River, van Donselaar 2548 (U); Maroni, 1901, Lem?e s.n.
(P). Para: Pararacweg at km 25 of road to Zanderij, Cramer LBB14902 (U). Saramacca: Zonneris
Creek near Bitagron, Cramer LBB14876 (U, VEN). Suriname: Lower Suriname R., Plantation Peper
pot, Cramer LBB14855 (U), Florsch?tz & Florsch?tz 1004 (U); Carolina, Middle Suriname R.,
Cramer LBB14856 (U); near Cassipora, Suriname R., Cramer LBB14857 (U); Domburg, Cramer
LBB14908 (NY, U); La Liberte, Beneden, Suriname R., Soeprato 179 (U). Unknown: valley of
Parepou Creek, Oldeman 2859 (U).?Trinidad. 1 mi S of St. Joseph, Hazen 2077 (GH, NY, US).?
Venezuela. Bolivar: Rio Uiri-yuk at El Foco, Maguire et al. 53507 (M, MY, VEN). Fed. Terr. Delta
Amacuro: Aniepo, Gines 4963 (US); Winiquina, Gines 5056 (US); Rio Amacuro, Sierra Imataca,
between Amacuro and mouth of Deadwater Creek Moat, Steyermark 87349 (F, NY).
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64 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 22. Nymphaea glandulifera (Hazen 2077). a. Undersurface of leaf. b. Opened flower, c.
Unopened flower bud. d. Upper surface of leaf.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 65
closure is completed by midnight, although flowers may remain open slightly longer
the first night. Only on the second night are the stamens exposed and, although not
so indicated, dehiscence probably occurs at this time, as in other members of subg.
Hydrocallis. The fruit production observed by Pagels in cultivation suggests that
self-pollination is likely in N. glandulifera and that the stigma remains receptive in
second-day flowers. Individuals of the scarab beetle Cydocephala castanea Olivier
were observed visiting flowers of N. glandulifera in Surinam.
The lack of evidence for stolon formation in N. glandulifera from herbarium
specimens or literature accounts implies that reproduction in this species occurs
entirely via seed production. The frequency with which maturing fruits are ob
served on herbarium specimens and are produced in cultivation indicates that
autogamy is probably involved in seed formation, as is the case in several other
species of subg. Hydrocallis.
On the basis of general morphology and floral biology, N. glandulifera ap
pears similar to N. conardii, N. gardneriana, and N. jamesoniana. These latter
three species normally exhibit a weblike leaf venation pattern that is not present
in N. glandulifera. Of the three, N. conardii is probably most closely related.
Both species have delicate leaves that are green above and below, lack stolons,
produce abundant seeds, and have very similar flowers. In addition to the differ
ence in leaf venation and some slight differences in leaf shape, the presence of
acicular sclereids in the stigmatic rays of N. conardii readily distinguishes it from
N. glandulifera. Also, the smooth seeds of N. glandulifera differ from the finely
granulate seeds of N. conardii. This feature can only be observed from seed testal
whole mounts with the aid of light microscopy. Certain features of N. glandulifera
seeds suggest an association with N. potamophila and N. oxypetala, two sagittate
leaved species. Further phytochemical and karyological study could provide addi
tional insight into this relationship.
The variation in pubescence which prompted the segregation of the glabrous
N. fenzliana, first as a distinct species and later as a variety of N. blanda, is not
considered to be worthy of taxonomic recognition. Similar variation in pubes
cence exists in N. tenerinervia, N. gardneriana, and N. amazonum.
The name Nymphaea blanda has been universally accepted as the correct
name of this species since its publication by Meyer in 1818; however, in publishing
N. blanda Meyer listed "N. glandulifera. Rodsch. Observ. p. 76" as a synonym
but rejected Rodschied's name as descriptively inappropriate. Meyer's citation
was a clear reference to Rodschied's Medizinische und chirurgische Bemerkungen
?ber das Klima, die Lebensweise und Krankheiten der Einwohner der hollaendi
schen Kolonie Rio Essequebo, where on page 76 N. glandulifera appears as
species number 89 with a short Latin description and some notes comparing it
with N. alba. Gerhard Wagenitz, who kindly supplied me with the relevant infor
mation from Rodschied's work in the University Library at GOET, agrees that N.
glandulifera was validly published by Rodschied. Therefore N. blanda must be
viewed as a superfluous name. Rodschied did not indicate a type specimen; his
collection (Rodschied 155) at GOET which is labelled N. glandulifera and con
forms to his description has been designated as the lectotype.
5. Nymphaea belophylla Trickett, Kew Bull. 26(1): 29. 1971.?Type: "a cultivated
specimen in the Kew Herbarium (K) grown at the Royal Botanic Gard
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66 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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X}m(>hata bclopkylla. i, leaf, X {; t,
SSIfeL* h gy:ioccium showing arrangement of
?wii, x 5. AH drawn from A. Blthtr. E
FIG. 23. Nymphaea belophylla. a. Photo of Trujillo 11430 (MY) from Barinas, Venezuela, b. Illustration repro
(Reproduced from Kew Bulletin, Vol. 26, by permission of the Controller of Her Majesty's Stationery Office.)
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68 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
empty folder at Kew is labelled N. belophylla; however, the staff was unable to
locate the type specimen or any surviving material of the original 1962 cultivation
at the Royal Botanic Gardens, Kew (reported as E.N. 167-62). No evidence
could be found of any specimen having been deposited at BM. The possibility of a
chance recovery of the type material and the absence of additional collections
which can be assigned with absolute certainty to this taxon make the designation
of a neotype inadvisable at this time.
Further collections from the region of the type locality are desperately needed
to permit a more accurate interpretation of this taxon. In the absence of this
material, the assessment of several critical features missing from the type descrip
tion has been dependent upon the assignment of Trujillo 11430 to N. belophylla.
This collection, which includes a leaf, flower, and attached tuber (Fig. 23a),
appears identical to N. belophylla as illustrated by Trickett (Fig. 23b). Certain
sagittate-leaved Nymphaea that have been collected in South America have car
pellary appendages shorter than those indicated for N. belophylla. These further
differ from the presumed TV. belophylla of Venezuela in lacking acicular sclereids
in the leaf and stigmatic ray tissues, where such structures are abundantly present
in the Venezuelan plants. The above differences form the basis for the segrega
tion of these plants as N. potamophila (q.v., no. 6).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 69
Specimens Examined. Brazil. Amazonas: Rio Negro, Padauiry, Telheiro, Fr?es 22686 (IAN);
Rio Negro, Paran? Ararira, ca. 100 km above Barcelos, Madison et al. 6155 (SEL, NY). Para: S?o
Francisco do Curur? Mission, boca de Igarap? Kenebiit-Taibiri para o Rio Curur?, Sioli 4 (IAN).
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70 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 25. Nymphaea potamophila (leaf from Madison et al. 6155; flower from Maia et al. 124). a.
Undersurface of leaf. b. Opened flower.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 71
acicular sclereids in the sepals. The floating sagittate leaves of N. potamophila are
reminiscent of N. oxypetala, which has sagittate submersed leaves, but floating
sagittate leaves of the latter species are reduced both in number and size. The
poorly known N. belophylla possesses sagittate floating leaves resembling those of
N. potamophila. Flowers of N. belophylla, as described by Trickett (1971), appear
to have carpellary appendages approaching twice the length of those in N. po
tamophylla. A plant assigned to N. belophylla, Trujillo 11430 (MY) from Barinas,
Venezuela, indeed displays these longer carpellary appendages. This plant is fur
ther distinguished in an abundance of acicular sclereids in the leaf and stigmatic
ray tissues, where these cells are completely absent in N. potamophila. If this
Venezuelan plant is in fact N. belophylla, the observed character differences
support its taxonomic separation from N. potamophila, in view of the conserva
tive nature of these characters among other species of subg. Hydrocallis. Accord
ingly, the two taxa have been segregated at the rank of species. However, addi
tional collection and study of these species in the South American interior are
clearly necessary to provide a better understanding of their relationship. Indeed,
the existence of a plant collected by Garc?a-Barriga (20813 at COL, GH) in
southeastern Colombia (state of Guainia) with leaves like N. potamophila but
flowers tending toward the presumed Venezuelan N. belophylla emphasizes this
need for further research.
7. Nymphaea oxypetala Planch?n, FI. Serres Jard. Eur. 8: 120. 1852. Nymphaea
raja Lehmann, Hamburger Garten- Blumenzeitung 9: 196, 208. 1853,
nom. superfl. Leuconymphaea oxypetala (Planch?n) Kuntze, Revis, gen.
pi. 1: 11. 1891.?Type: Ecuador, "in inundatis circa Guayaquil," Mar
1845, Jameson 334 (holotype: G!; isotypes: B! BM! K!). The specimen at
B (from the herb. Lehmann) contains label of Parker from Guyana and
no information of Jameson (see discussion).
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72 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
suffused with purple, containing acicular sclereids, splitting along three longitudi
nal sutures adjacent to anther sacs to release tiny spherical "sclerites," the
outermost to 6.5 cm long, with filaments to 16 mm wide and connectives pro
longed 1.5 mm or more beyond anthers, the inner with filaments 1-2.4 mm wide
and connectives prolonged to 1 mm beyond anthers, the inner anthers 0.58-0.73
times as long as stamen; pollen diameter ca. 40 \im. Carpels 21-27, the append
ages 13-29 mm long, cream-colored, abruptly bent at distal %, linear-clavate, with
enlarged tips 6-20 mm long, 1-2 mm in diameter, the stigmatic rays shortened
axially, lacking acicular sclereids, with papillae cells remaining attached; axial
process absent. Fruits normally present; seeds 0.85-1 mm long, 0.7-0.85 mm in
diameter, 1.1-1.25 times as long as broad, with clusters of hairs 70-120 [xm long,
the fine surface topography smooth. Tuberiferous flowers absent. Chromosome
number: 2n = 84. Figs. 1, 6, 10, 12, 26.
Phenology. Flowers have been collected during March in Ecuador, August in
Venezuela, and April and July in southern Brazil; fruits were obtained during
August in Venezuela and July in Brazil.
Distribution. (Fig. 24). Collected in Ecuador, Venezuela, and southern
Brazil. Material of N. oxyp?tala is also present in mixed collections of Nymphaea
from Cuba and Paraguay, but until additional collections from both areas are
obtained the accuracy of these records is questionable. Attempts at cultivation of
N. oxypetala (from Wiersema & Gonz?lez 2232) indicate a low level of tolerance
to increased turbidity or salinity and decreased temperatures. Although the small
number of collections of N. oxypetala provides little ecological information, this
submersed species is probably best adapted to clean, gently flowing freshwater
habitats.
Specimens Examined. Cuba. Habana: Hoyo Colorado, brook near Laguna Ariguanabo, Ekman
13710 (S, flower only, leaf is N. amazonum).-Brazil. Mato Grosso: Biarra do Rio dos Bugres,
Lindman 2877 (S). Mato Grosso do Sul: Corumba, Malme 1764B (S).?Ecuador. Prov. unknown: in
num. Daule, Mille 1000 (F).?Paraguay. Gran Chaco: Santa Elisa, Hassler 2804 (G, K, both with
flower of N. oxypetala mounted with material of N. prolif?ra).?Venezuela. Apure: near San Fer
nando de Apure, Ramia & Montes 5362 (VEN); Guasdualito, Fern?ndez 1299 (MY), Wiersema &
Gonz?lez 2232 (F, MO, MY, NY, VEN, UNA). Cojedes: R?o Portuguesa, Delascio et al. 4493
(VEN).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 73
FIG. 26. Nymphaea oxypetala (Wiersema & Gonz?lez 2232). a. Undersurface of subm
b. Upper surface of floating leaf. c. Opened flower, d. Cross section of petiole, e. Cro
peduncle.
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74 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
oxypetala. His citation of type material reads: "in Columbia collegit Cl. Jameson,
in Chile filius Guilielmus Lehmann." The name N. raja, handwritten by Leh
mann, appears on one of two sheets of the 1845 Jameson collection from Guay
aquil (until 1830 included in Great Columbia) at K and on a specimen from
Lehmann's own herbarium (now at B). This latter specimen bears no information
of Jameson and is now labelled "Mr. Parker, Guiana." Caspary (1878), who
viewed this specimen, makes no mention of the Parker label. He does indicate,
however, that he was unable to find evidence of a collection from Chile by
Lehmann's son either in Lehmann's herbarium or in any other herbarium. As
Parker's collections from Guyana in other herbaria have never included N. oxype
tala (which has never been found in Guyana) and the Jameson collection of this
species is the only one prior to 1890 in existence, Lehmann's specimen is undoubt
edly material collected by Jameson in Ecuador.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 75
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76 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Specimens Examined: Colombia. Bolivar: Soplaviento and vicinity, Killip & Smith 14613 (F, GH,
NY, US). Magdalena: Isla de Salamanca, Los Almendros, Romero-Casta?eda 10869 (COL).?
Venezuela. Falc?n: between Chimpire & El Guay, road Coro to Tucacas, Prance 23722 (NY, UNA);
vicinity of Tucacas, Prance 23724 (NY, UNA); Paraguana, Tamayo 701 (VEN); 1.5-2 km N of
Tucacas, Steyermark & Manara 110570 (NY, VEN); near dam of Jatira, environs of Tucuyo de la
Costa, Trujillo 13467 (F, MY). Gu?rico: environs of El Sombrero, Trujillo 2538 (MY); km 2 between
crossing at Barbacoas and crossing at El Sombrero, Trujillo 11329 (MY); km 10 from the crossing La
Guamita (El Sombrero-Chaguaramas) and Barbacoas toward Barbacoas, Trujillo 11358 (MY); off hwy
2, ca. 2 km S of R?o Gu?rico near Calabozo, Wiersema 1404 (VEN); 25 km S Calabozo, Wiersema
1411 (NY, UNA, US, VEN); 34.8 km S of R?o Gu?rico at Calabozo, Wiersema 1420 (BM, COL, MO,
UNA, VEN).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 77
stamens dehisce in the early evening, they are not exposed until ca. 1-2 a.m. and
continue spreading thereafter. Second-day flowers close at dawn and begin to be
retracted under water. Flowers both nights are only faintly odorous. Although the
stigma is fairly dry by the second night, it is still receptive to pollen, as some
flowers in cultivation produced seeds autogamously. This autogamous seed pro
duction was observed in only a few flowers, however, with the majority of flowers
failing to set seed in the absence of cross-pollination.
No evidence of asexual reproduction exists for N. novogranatensis, which
apparently relies entirely on seed production for its dispersion. The data obtained
indicate that this species is less efficient in autogamous seed production than other
more widespread species of subg. Hydrocallis, such as N. jamesoniana or N.
conardii. This factor may be partly responsible for the restricted distribution of TV.
novogranatensis, which is thus more dependent on pollinating organisms for suc
cessful reproduction.
Nymphaea novogranatensis is most closely related to N. tenerinervia, the two
having almost identical floral morphology and biology. In addition to the differ
ences presented in the key, N. novogranatensis differs in having glabrous petioles
and lacking stolons (vs. glabrate or pubescent petioles and stolon formation in N.
tenerinervia). The mucronulate leaf apices of N. novogranatensis serve to distin
guish it from all other members of the subgenus, which have rounded or truncate
leaf apices.
9. Nymphaea tenerinervia Caspary in Martius, Fl. bras. 4(2): pi. 36. 1878.?Type:
Brazil. Bahia: "prope Juazeiro ad flumen St. Francisco," April 1819,
Martius s.n. (lectotype here designated: M!, mounted on 2 sheets; isolec
totype: B?, see discussion).
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78 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Specimens Examined. Brazil. Cear?: BR-222 at km 16,16 mi W Fortaleza, Wiersema et al. 2314
(GUA, IPA, MO, UNA). Goi?s: Brejinho de Nazare, entre Fatima e Alian?a do Norte, 2 km N de
Rio Crixas, Camargo de Abreu 25 (NY, SP). Maranh?o: BR-316 at km 598 between Teresina and
Caxias, Wiersema et al. 2307 (INPA, IPA, UB, UNA). Para: 12 km de Bragan?a, Maciel & Bou?as 43
(IAN). Piaui: BR-343 Teresina-Sobral at km 303, Rio Suruben, Wiersema et al. 2310 (IPA, NY, SP,
UNA); BR-343 at km 260 E of Campo Meior, Rio Jenipapo, Wiersema et al. 2311 (IPA, UNA).
Roraima: Rio Branco, Surumu, Ule 8120 (G, K, not B).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 79
FIG. 29. Nymphaea tenerinervia (Wiersema et al. 2314). a. Undersurface of leaf. b. Opened
flower, c. Unopened flower bud. d. Cross section of petiole, e. Cross section of peduncle, f. Upper
surface of leaf.
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80 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
10. Nymphaea amazonum Martius & Zuccarini, Abh. Math.-Phys. Cl. K?nigl.
Bayer. Akad. Wiss. 1: 363. 1832. Nymphaea blanda ? amazonum (Mar
tius & Zuccarini) Planch?n, FI. Serres Jard. Eur. 11: 21. 1856. Nymphaea
rudgeana ? amazonum (Martius & Zuccarini) Grisebach, Fl. Brit. W.
Ind. 12. 1864. Leuconymphaea amazonum (Martius & Zuccarini) Kuntze,
Revis, gen. pi. 1: 11. 1891. Castalia amazonum (Martius & Zuccarini)
Britton & P. Wilson, Bot. Porto Rico 5(2): 305. 1924.?Type: Brazil.
Para: "in aquis stagnantibus prope urbem Para," May-June (probably
1819), Martius exsicc. 3313 (holotype: M!, mounted on 3 sheets).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 81
1. Leaf blades 1-1.35 times as long as wide, with trichosclereids dense on upper surface,
acicular sclereids absent from mesophyll; stamens mostly less than 200.
10a. N. amazonum subsp. amazonum.
1. Leaf blades 1.25-1.5 times as long as wide, with trichosclereids sparse on upper surface,
acicular sclereids often present in mesophyll; stamens usually more than 225.
10b. N. amazonum subsp. pedersenii.
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82 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Representative Specimens. Cuba. Habana: Guatao, Le?n 11522 (NY); Laguna de Perdig?n,
Coralillo, Le?n & Nivaro 7609 (NY); Hoyo Colorado, near Laguna Ariguanabo, Ekman 13710 (S).?
Dominican Republic. Saman?: Peninsula de Saman?, S?nchez, Ekman 15949 (G, S).?Guadeloupe.
Pointe Noire, Duss 3702 (F, GH, MO, NY, US); Vieux-Bourg, Raynal-Roques & J?r?mie 21101
(P).?Haiti. Dept. du Nord, Bayeux, between Bord de la Mer and Morne ? Madame, Ekman 2680
(S).?Jamaica. Locality unknown, March 25 (GH, K).?Martinique. Le Marin, entre Cap Ferr? et
Barri?re la Croix, Raynal-Roques & J?r?mie 21169 (P).?Puerto Rico. Punta Santiago, Stevens 2463
(NY); Sauturee, Britton & Britton 7086 (NY); Sabana Atiajo, Britton & Britton 9234 (NY); Vega
Baja, Britton et al. 6762 (F, NY, US).
Mexico. Tabasco: Laguna del Corcho, near Ocuiltzapotl?n, Rovirosa 674 (NY, PH, US).
El Salvador. Laguna de Zapotitan, Fassen 29326 (WIS).?Guatemala. Peten: near Flores,
Hedger 75 (BM).?Nicaragua. Chontales: road from Acoyapa to San Carlos, 2 km NW of Rio Oyate,
Haynes 8593 (MO, UNA).?Panama. Canal Zone: Col?n, Lehmann 1007 (BM). Panam?: road be
tween Panam? and Chepo, Dodge et al. 16713 (MO).
Bolivia. Santa Cruz: Santa Cruz, Kuntze 340 (NY).?Brazil. Acre: 4 km from Brasileia, Lowrie
et al. 720 (INPA). Alagoas: BR-316, 1 km E Maribondo, Wiersema et al. 2291 (IPA, UNA). Bahia:
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 83
FIG. 30. Nymphaea amazonum. a-f. N. amazonum subsp. amazonum (Wiersema et al. 2304). a.
Undersurface of leaf. b. Opened flower, c. Unopened flower bud. d. Cross section of petiole, e. Cross
section of peduncle, f. Upper surface of leaf, g, h. N. amazonum subsp. pedersenii (Wiersema et al.
2270). g. Undersurface of leaf. h. Upper surface of leaf.
Lac da Con?eic?o, Blanchet 11 (G); 10 km NE Salvador, Lagoa de Abaete, Anderson 224 (INPA).
Cear?: Mun. Fortaleza, Barra do Cear?, Drouet 2511 (F, GH); km 204 of BR-222 between Sobral
and Fortaleza, Wiersema et al. 2312 (IPA, UNA). Maranh?o: BR-316 at km 613, 8 km W of Rio
Parnaiba, Wiersema et al. 2304 (IPA, UNA). Mato Grosso: Mun. Pocon?, km 62 Rodovia Transpanta
neira, Rio Pixaim, Macedo et al. 452 (INPA), km 63, da Silva 452 (UFMT). Minas Gerais: Mun. Santa
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84 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 31. Distribution of Nymphaea amazonum subsp. amazonum and subsp. pedersenii.
Luzia, Lagoa Santa, Warming 1750 (C), Barreto 7230 (BHMH). Para: Mun. Soure, Porto Santo
Antonio, Black 50-9086 (IAN); Rio Cupari, Fl?chai, Black 47-2105 (IAN). Pernambuco: Dois Ir
mois, near Recife, Wiersema et al. 2284 (IPA, UNA). Piaui: Lake at Parnagua, Gardner 2474 (BM);
BR-316 at km 240 between Picos and Teresina, Wiersema et al. 2303 (IPA, UNA). Rio de Janeiro:
Mun. Cabo Fr?o, Pontal Beach, Smith et al. 6647 (NY, US), Restinga 1307 (NY); Mun. S?o Jo?o da
Barra, between S?o Jo?o da Barra and Atafona, Restinga 418 (US); Mun. Campos, Lagoa Feia,
Araujo & Maciel 3822 (GUA). Rond?nia: Rio Guapor?, Faz. Cruz Verde, Black & Cordeiro 52
15053 (IAN).?Colombia. Cauca: Locality unknown, Holton 689 (G, GH, NY, PH); Valles du Cauca,
1851-1857, Triana s.n. (BM, K, P). Magdalena: Valles de Magdalena, Triana 5020 (COL). Tolima:
llanos de Ibaqu?, 1851-1857, Triana s.n. (BM). Valle: entre Jamundi y R?o Pance, Cuatrecasas 19656
(COL, F, G); km 31 of hwy Cali-Popayan, Maas & Plowman 2159 (COL, U, VEN) .?Ecuador.
Guayas: hwy 25, 21 km S Jujan, Wiersema et al. 2027 (UNA, USF); hwy 25, 8 km N Guayaquil,
Wiersema et al. 2029 (AAU, F, NY, UNA, US, USF). Morona-Santiago: Lago de Kumpaka, 12 Aug
1982, Steinitz-Kannan s.n. (UNA).?French Guiana. Sinnamary, Raynal-Roques 19861 (P); Kaw, 60
km SE de Cayenne, Raynal-Roques 21545 (P).?Guyana. Coast Canals, Jenman 5747 (BM, K, NY,
U); dam road from Mon Repos to Water Conservancy, Cramer 18 (U); Georgetown, Cramer 23 (U);
Berbice River, new Dageraad, Maas et al. 5565 (UNA).?Peru. Loreto: Balsopuerto, Killip & Smith
28682 (NY, US). Madre de Dios: 30 km SSW Puerto Maldonado, Barbour 4885 (MO, UNA).?.
Surinam. Commewijne: Plantation Koonenburg, Lower Commewijne River, Cramer LBB14849 (U).
Coronie: Inverness, km 129.5, Cramer LBB14851 (COL, U). Nickerie: Prins Bernard-polder, Hek
king 940 (C, GH, K, NY, U, VEN). Para: km 24 of road to Zanderij, Cramer LBB14846 (U).
Saramacca: Zonneris Creek near Bitagron, Cramer LBB14877 (F, U). Suriname: Weg naar Zei, NW
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 85
of Paramaribo, Cramer LBB14847 (U).?Tobago. The Whein, Broadway 3366 (BM, F, MO, NY,
PH); Studley Park, Broadway 4560 (BM, F, G, GH, K, NY, U, US).?Venezuela. Apure: Sabanas
de Arichuna, Ramia & Montes 5532 (VEN). Aragua: Dist. Urdaneta, 10 km SE Camatagua, Wier
sema 1436 (NY, UNA, VEN). Delta Amacuro: near Tucupita, Trujillo 12860 (MY). Gu?rico: Dist.
Miranda, 12 km N Calabozo, Wiersema 1400 (UNA, US, VEN). Miranda: Dist. Bri?n, sabana de
Guardalagua E of Sotillo, Wiersema & Gonz?lez 2203 (F, MY, NY, UNA, VEN). Monagas: Isla de
Guara, Depto. Sotillo, Mpio Uracoa, Trujillo 12946 (MY). Sucre: Laguna de Campoma, Mpio Cari
aco, Trujillo 11917 (MY); Trujillo 14380 (MY).
Flowers of this subspecies are floating and open two successive nights (Con
ard 1905; W. Pagels, pers. comm.; Wiersema, unpubl.). First-day flowers remain
fully closed until 2-3 hours before dawn. The sepals and outer petals then open
very rapidly to provide an aperture ca. 1 cm in diameter to the center of the
flower about one hour before dawn. The stigmatic surface is covered with fluid
and with the powdery remains of papillae cells. Closure of the flower is very rapid
and completed by dawn. Second-day flowers begin opening at dusk but become
fully open only about one hour before dawn. All the stamens are now fully
exposed, their anthers dehiscing at this time. The carpellary appendages are in
curved over the stigmatic surface, which is now dry. Closure of second-day
flowers begins at dawn and is completed in about one hour, the flower being
slowly retracted under water. Flowers both nights emit a strong aromatic odor
that has been variously described as resembling turpentine, xylol, petrol, ben
zene, xylene, PDB, benzol, or acetone. In Surinam they are reportedly visited by
the scarab beetle Cyclocephala verticalis Burmeister (Cramer et al. 1975). From
observations of plants in cultivation (from Wiersema 1400), it is evident that seeds
are produced autogamously. The stigma therefore remains receptive in second
day flowers.
Nymphaea amazonum subsp. amazonum relies mainly on autogamous seed
production for dispersion. Plants also propagate by the formation of elongate
stolons or short sprouts from the rhizome throughout the growing season.
A sterile Brazilian population in southwestern Pernambuco (Wiersema et al.
2296 at IP A, UNA) appears, on the basis of morphological and chemical evi
dence, to represent a natural hybrid between N. amazonum subsp. amazonum
and N. lasiophylla. Other plants collected from the nearby state of Minas Gerais
(Mos?n 803, Regnell 2036, both at S) may also represent hybrids involving N.
amazonum subsp. amazonum and TV. lasiophylla.
Several collections from central Colombia display a ring of pubescence at the
peduncular apex in addition to that generally found at the petiolar apex. In the
absence of other detectable morphological difference, no special taxonomic status
is being assigned to these plants. They have previously been regarded as a distinct
species, N. goudotiana, or as a variety or form of N. amazonum. The variety
forma-submersa reported by Sagot (1881) from French Guiana is actually N.
rudgeana.
Lehmann (1853) based his N. candolleana on N. ampia sensu de Candolle,
but explicitly excluded the basionym Castalia ampia Salisbury from his taxon. In a
footnote he mentioned a specimen of Splitgerber from Surinam as being the most
representative of his species. Presumably the lectotype at B is this specimen,
which is clearly N. amazonum subsp. amazonum.
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86 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Specimens Examined. Argentina. Corrientes: Depto Mburucuy?, Estancia Santa Teresa, Peder
sen 4494 (C, CORD, K, NY, P, S), Schinini & Crist?bal 10791 (CTES); Depto Mercedes, Rio
Mirinay, Krapovickas & Crist?bal 21733 (CTES, G, MO, WIS), Colonia Pellegrini, Quarin & Schinini
1063 (CTES); Depto Monte Caseros, Monte Caseros, arroyo Timboa, Nicora 7132 (CTES); Ruta 127
and arroyo Curuz? Cuatia, Schinini et al. 17395 (CTES); Depto San Mart?n, Colonia Pellegrini,
Laguna Ibera, Krapovickas et al. 29505 (CTES).?Brazil. Mato Grosso: Pocon?, Macedo &
Assumps?o 688 (NY); Transpantaneira Highway, Fazenda Jofre, Prance et al. 26205 (NY), da Silva
395 (UFMT); Rodavia Transpantaneira km 20, da Silva 440, 441 (UFMT), km 22, da Silva 459
(UFMT); Mun. C?ceres, Ilha Taiama, da Silva 212 (UFMT).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 87
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88 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
connectives prolonged 0.5-2 mm or more beyond anthers, the inner with fila
ments to 1 mm wide and connectives prolonged to 0.6 mm beyond anthers, the
inner anthers 0.6-0.85 times as long as stamen; pollen diameter 30-40 |im. Car
pels 20-39, the appendages 6-15 mm long, indigo or purple, with cl?vate tips
1.5-4 mm long, 0.6-1.9 mm in diameter, the stigmatic rays lacking acicular
sclereids and with papillae cells separating into powdery mass; axial process ca. 1
mm long, acute to obtuse. Fruits and seeds rarely if ever produced. Seeds from
artificial hybridization finely granulate, with longitudinal rows of hairs. Tuberifer
ous flowers usually present, submersed; outer sepaloid bracts (3-) 4, 3-6 cm long,
1.8-2.3 cm wide, greenish to reddish, usually with short black streaks, subcoria
ceous, ovate-triangular with truncate base; tuber globose, densely wooly, produc
ing alternating whorls of tuberous flowers and leaves in (3's) 4's; peduncle of
tuberiferous flowers to 7.5 mm in diameter, glabrous, abcission layer forming
near apex. Chromosome number: 2n = 18. Figs. 10, 12, 32.
Phenology. Collected in flower from November through June, although
flowering appears to be of shorter duration further south; no collections of fruits
from natural populations are known.
Distribution. (Fig. 28). Collected only in northern Argentina, Paraguay,
southern Brazil, western Ecuador, and in Costa Rica and El Salvador; primarily
in lowland savannahs.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 89
FIG. 32. Nymphaea prolif?ra (Wiersema & Pedersen 2248). a. Undersurface of leaf. b. Upper
surface of young leaf. c. Opened flower, d. Cross section of petiole, e. Cross section of peduncle, f.
Upper surface of leaf. g. Tuberiferous flower, h. Cross section of peduncle of tuberiferous flower.
before dawn. Autogamous seed production has not been observed in N. prolif?ra,
which suggests that the stigma is no longer receptive in second-day flowers.
Although seeds have been produced from an artificial hybridization in cultiva
tion, no evidence of seed production has been observed in field populations. This
is apparently due to an absence of available pollinators in many of the regions N.
prolif?ra now inhabits. In northern Argentina, the subtemperate nature of the
climate may exclude potential pollinators of N. prolif?ra from tropical regions to
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90 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
12. Nymphaea lasiophylla Martius & Zuccarini, Abh. Math.-Phys. Cl. K?nigl.
Bayer. Akad. Wiss. 1: 364. 1832, quoad folium, flos est N. tenerinervia
Caspary. Leuconymphaea lasiophylla (Martius & Zuccarini) Kuntze, Re
vis, gen. pi. 1: 11. 1891.?Type: Brazil. Bahia: "in aquis stagnantibus ad
flumen St. Francisci prope Juazeiro," April-May (probably 1819), Mar
tius exsicc. 2377 (lectotype, the leaves only designated: M!, on 2 sheets;
isolectotype: B!-fragment).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 91
diameter, glabrous, with 6 or 18 major air canals. Flowers floating, the sepals and
outer petals in distinct whorls of four. Sepals 3.5-7 cm long, 1.3-3.3 cm wide,
uniformly green or yellow-green, ovate, acute or somewhat rounded at apex,
lacking acicular sclereids, inconspicuously nerved. Petals 20-26, creamy white
with lace work venation, lacking acicular sclereids, the outer only slightly shorter
than sepals, with apex rounded to somewhat acute, the inner 3.1-5.9 cm long; 1
2.5 cm wide, acute or somewhat rounded; transition to stamens abrupt. Stamens
65-130, creamy white, containing fine acicular sclereids, the outermost to 4.7 cm
long, with filaments to 9 mm wide and connectives prolonged 0.5-1.5 mm beyond
anthers, the inner with filaments to 1.8 mm wide and connectives prolonged 0.2
mm or less beyond anthers, the inner anthers 0.4-0.6 (-0.7) times as long as
stamen; pollen diameter 35-40 ^m. Carpels 19-33, the appendages 5-14 mm
long, to 3 mm wide near base, with reddish purple tips, lingulate-tapering, the
stigmatic rays lacking acicular sclereids and with papillae cells remaining attached;
axial process minute or absent. Fruits seldom develop; seeds 0.9-1.05 mm long,
0.6-0.7 mm in diameter, 1.4-1.6 times as long as broad, with longitudinal rows
of hairs 55-110 \xm long, the surface finely ruminate. Tuberiferous flowers usually
present, submersed; outer sepaloid bracts 3-4, uniformly yellow-green or green;
inner bracts 0-4; tuber globose, densely wooly, usually producing alternating
whorls of tuberous flowers and leaves in 3's; peduncle of tuberiferous flower to 7
mm in diameter, glabrous, abscission layer forming near apex. Chromosome num
ber: 2n = 18. Figs. 3, 10, 12, 33.
Phenology. Collected in flower throughout the year; fruiting collections are
rare, in June and August.
Distribution. (Fig. 34). Known only from coastal states of eastern Brazil, Rio
de Janeiro north to Piaui, with a disjunct population, probably introduced, on Isla
de Margarita, Venezuela; usually in stagnant water, often in artificial ponds.
Specimens Examined. Brazil. Alagoas: between Palmeira dos Indios and Bom Conselho, 12 km
S of division Alagoas-Pernambuco, Wiersema et al. 2293 (IPA, MO, UNA). Bahia: E of Rio Salitre,
between Juazeiro and Sobradinho, Wiersema et al. 2298 (IPA, UNA). Cear?: Mun. Fortaleza, Lagoa
Mecejana, Mecejana, Drouet 2140 (F, GH, S, US), Lagoa do Tanape, Drouet 2292 (F, GH, US),
Drouet2563 (GH), ponds of Damas, Drouet 2486 (F, GH, S, US); Tat?, L?fgren 729 (S); km 195 of
BR-222 between Sobral and Fortaleza, Wiersema et al. 2313 (INPA, IPA, UNA, US); km 14 of BR
304 between Fortaleza and Mossor?, Wiersema et al. 2316 (IPA, MO, SP, UNA, US). Paraiba: a?ude
Novo, Campina Grande, Wright 1 (GH); a?ude Puxanana, Wright 4 (F, GH). Pernambuco: in a river,
Tapera, Pickel 3725 (F, GH, IPA, NY); km 327-8 of BR-232 between Arcoverde and serre Talhada,
Wiersema et al. 2294 (IPA, UNA); km 475 of BR-232 E of Salqueiro, Wiersema et al. 2295 (IPA, NY,
?B, UNA); Mun. Rajada, km 45 of PE-407 between Petrolina and Paulistana, Wiersema etal. 2302.
Rio de Janeiro: Mun. Campos, BR-356, 1 km de Campos, Casari et al. 523 (GUA). Rio Grande do
Norte: Mossor?, Wiersema etal. 2317 (F, GUA, IPA, MO, UNA).?Venezuela. Nueva Esparta: Isla
de Margarita, between Boca del R?o and Boca del Pozo, Steyermark et al. 130978 (MO, UNA).
The floral behavior has not been studied; however, flowers open fully when
placed in darkness and emit a strong aromatic odor.
In northeastern Brazil reproduction in N. lasiophylla is primarily asexual.
Propagation in most populations is by means of abortive tuberiferous flowers, a
phenomenon among Nymphaea common only in this species and N. prolif?ra,
although rare occurrences of similar structures have been reported for other syn
carpous Nymphaea (Mohan Ram & Nayyar 1974; Mitra & Subramanyam 1982).
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92 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 33. Nymphaea lasiophylla (leaf from Wiersema et al. 2317; flower from Wiersema et al.
2316). a. Undersurface of leaf. b. Opened flower, c. Cross section of petiole, d. Cross section of
peduncle, e. Upper surface of leaf.
These abortive flowers, their status indicated by the peduncular air canal anatomy
and the production of outer sepaloid bracts, develop an enlarged tuber, which
produces whorls of leaves and additional secondary tuberiferous flowers. A few
orders of branching may result from similar development of secondary and even
tertiary tuberiferous flowers, resulting in the production of a substantial number
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 93
of tubers. The tubers detach readily, remain afloat briefly, and eventually develop
into adult plants. Leaves and roots are often well-developed prior to abscission of
these tubers, which suggests the application of the term vivipary to this reproduc
tive strategy. The ratio of tuberous to normal flowers varies from one population
to another. Although tuberous flowers were present in all field populations
studied, normal flowers were not observed in some populations. Another form of
vegetative propagation, the production of stolons, has been observed in certain
populations as well.
Seed production was evident in only one of ca. 20 field populations examined.
Coincidentally, in only this and another nearby population (ca. 1 km away) was
there evidence of pollinating beetles, these found trapped within some of the
diurnally closed flowers. This suggests that cross-pollination may be necessary for
seed production, the absence of seeds from most populations being attributable to
an absence of pollinators. Further study is necessary to verify this hypothesis.
Nymphaea lasiophylla is clearly linked to N. lingulata; the two species are
unique among members of subg. Hydrocallis in their possession of tapering (as
opposed to cl?vate) carpellary appendages. Karyology, phytochemistry, and seed
morphology all support this close relationship. Although not apparent phyto
chemically, N. lasiophylla must also bear some relation to N. prolif?ra, as both
species exhibit a form of vegetative reproduction not observed elsewhere in the
subgenus. A putative hybrid involving N. lasiophylla and N. amazonum subsp.
amazonum is discussed under this subspecies (no. 10a).
The type description, drawn from Martius's field notes and the type speci
mens, is based on a mixed collection (gathered in 1819 fide Caspary, 1878). The
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94 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 95
FIG. 35. Nymphaea lingulata (Wiersema et al. 2308). a. Undersurface of leaf. b. Opened flower,
c. Unopened flower bud. d. Cross section of petiole, e. Cross section of peduncle, f. Upper surface of
leaf.
anthers, the inner anthers 0.7-0.8 times as long as stamen; pollen diameter 30-60
pm. Carpels 20-28, the appendages 5-14 mm long, to 2 mm wide at base, creamy
white with purplish base abaxially, lingulate-tapering, the stigmatic rays lacking
acicular sclereids and with papillae cells remaining attached; axial process ca. 1
mm long and blunt. Fruits known only from type collection; seeds 1.05-1.2 mm
long, 0.8-0.9 mm in diameter, ca. 1.3 times as long as broad, with longitudinal
rows of short hairs 30-70 \xm long, the surface finely ruminate. Tuberiferous
flowers absent. Chromosome number: 2n = 18. Figs. 3, 10, 12, 13, 35.
Phenology. Flowering material scanty, collected in February, April, June, and
August; fruits known only from my own June collection.
Distribution. (Fig. 34). Collected only in northeastern Brazil where observa
tions indicate it is probably quite rare. Collections from Paraguay which were
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96 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
previously assigned to this species (Wiersema 1984a) are now believed to repre
sent N. gardneriana.
Specimens Examined: Brazil. Cear?: Mun. Maranguape, a?ude S?o Bento, Drouet 2174 (F,
GH). Minas Gerais: Mun. Pirapora, Alagoas, Barreto 11319 (BHMH, UNA); Mun. Ituiutaba, L.
Teyiuba, Macedo 2279 (MO, S); Lagoa da Babylonia, Glaziou 12414 (C, G, K, P).
14. Nymphaea rudgeana G. Meyer, Prim. fl. esseq. 198. 1818. Nymphaea ampia ?
rudgeana (G. Meyer) de Candolle, Syst. nat. 2: 54. 1821. Castalia rudge
ana (G. Meyer) Britton & P. Wilson, Bot. Porto Rico 5(2): 305. 1924.?
Type: Guyana. "In rivulis continentis," Rodschied 281 (lectotype, desig
nated by Caspary, 1878: GOET!). The type sheet contains one leaf and
one flower of N. rudgeana and one flower of N. glandulifera, the latter
undoubtedly from Rodschied 155, which has apparently been mismounted
from the original specimen of the Grisebach Herbarium.
Nymphaea sinuata Salzmann ex Lehmann, Hamburger Garten- Blumenzei
tung 9: 203. 1853.?Type: Brazil. Para: Lago d'Obidos, Spruce 479 (lec
totype, here designated: K!).
Nymphaea amazonum var. forma-submersa Sagot, Ann. Sei. Nat. Bot., ser.
6, 2: 142. 1881.?Type: French Guiana. Karouany [Acarouani], 1855,
Sagot 25 (lectotype, here designated: K!).
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 97
Representative Specimens. Cuba. Santa Clara: city of Santa Clara, Britton etal. 6129 (NY, US);
near Lomo Cruz, Britton et al. 10238 (NY).?Guadeloupe. Locality unknown, Duchassaing 16
(GOET).?Jamaica. St. Elizabeth: Black River, Adams 13218 (UCWI).?Martinique. Le Robert,
NE de Trou Terre, Raynal-Roques & J?r?mie 21178 (P).
Nicaragua. San Juan del Norte (Greytown), Stevens 20848 (MO).
Brazil. Amap?: Fazenda Mungubas, Black & Lobato 50-9534 (IAN); lago grande, Goeldi 1204
(BM, INPA, U). Amazonas: Taperinha bei Santar?m, Ginzberger & Hagmann 658 (F, W); Mun.
Humayta, near Livramento, Krukoff6969 (BM, BR, F, GH, K, LIL, MO, NY, U, US). Bahia: 5 km S
of Santa Cruz, Harley et al. 17129 (MO, NY, P, U); 10 km NE of Salvador, Anderson 223 (INPA); Rio
Salitre, between Juazeiro and Sobradinho, Wiersema etal. 2299 (IPA, UNA). Cear?: Mun. Fortaleza,
Barra do Cear?, Drouet 2512 (F, GH, S, US). Maranh?o: S?o Luis, Anderson 161 (NY, INPA); km 583
of BR-316, 38 km W Rio Parnaiba, Wiersema et al. 2309 (IPA, UNA). Para: lina do Mosquiero, near
Para, Killip & Smith 30531 (NY, US); Mun. Bel?m, Lagoa Agua Pr?ta (Utinga), Drouet 1955 (F, GH,
US); Rio Arapiuns, Lago Mentai, Pires & Silva 4359 (IAN, US). Paran?: Mun. Paranagua, Pontal do
Sul, Hatschbach 16387 (US). Pernambuco: Tapera, Pickel 2343 (F, GH, US); Dois Irmois near Recife,
Wiersema et al. 2285 (IPA, ?B, UNA); km 45 of PE-60 S of Cabo, Wiersema et al. 2286 (IPA, NY,
UNA). Rio de Janeiro: Mun. Campos, Farol de S?o Tom?, Araujo & Maciel 4056 (GUA); Mun. de
Casimiro de Abreu, Lagoa Iodada, Casari 500 (GUA). S?o Paulo: Santos, Mos?n 3339 (BR, C, GH,
GOET, K, M, S, U).?Colombia. Magdalena: Santa Marta, Smith 2108 (BM, BR, G, GH, K, NY, P,
PH, US).?French Guiana. Vicinity of Cayenne, Broadway 276 (GH, NY, US); Cayenne, Badocul,
Broadway 919 (GH, NY, US); Savane Gabrielle, 10 km ESE de Stoupan, Raynal-Roques & J?r?mie
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98 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
FIG. 36. Nymphaea rudgeana (Wiersema & Gonz?lez 2205). a. Undersurface of leaf. b. Opened
flower, c. Unopened flower bud. d. Cross section of petiole, e. Cross section of peduncle, f. Upper
surface of leaf.
21291 (P); Kaw, 60 km SE de Cayenne, Raynal-Roques 21554 (P).?Guyana. Demerara, 1824, Parker
s.n. (K, NY); Barina River, Jenman 7230 (K); Kara-Kara Creek, Wismar, Demarara River, Jenman
7779 (K); Tapacooma Lake, Jenman 7892 (K); Kaieteur Plateau, Cowan & Soderstrom 2224 (K, NY,
US); Amacuro River, NW District, de la Cruz 3563 (GH, US); Wanama River, NW District, de la Cruz
3969 (F, GH, MO, NY, PH, US); Rapununi, Moreru Lake, Goodland 1069 (US); Lamaha stop-off, SE
of Georgetown, Hitchcock 16868 (NY, S, US).?Surinam. Brokopondo: Kwakoegron, Saramacca
River, Maguire 25004 (F, GH, K, NY, US). Commewijne: Voorburg, Lower Suriname River, Cramer
LBB14861 (U). Coronie: Sarah-Leasowes, Cramer LBB14865 (MO, U). Marowijne: Maengo, Cramer
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 99
LBB14913 (F, U). Nickerie: Nieuw-Nickerie, Hekking 955 (GH, K, NY, U). Para: Coropina Creek,
Cramer LBB14859 (COL, NY, U). Suriname: 2 km E of Meerzorg, Cramer LBB14860 (U).?Trinidad.
1 mi S St. Joseph, Hazen 2079 (NY); S of Arima, near Caroni River, Britton & Britton 2887 (GH, K,
NY, US).?Venezuela. Anzoategui: R?o Tigre, ESE of San Tom?, Pittier 1408 (VEN); km 70-71 de
carretera Sn. Mateo-Anaco, Rutkis 169 (VEN). Bol?var: near Urull?n, at foot of Auyantepui, Vareschi
& Foldats 4532 (NY, VEN); drainage of R?o Aponguao, 152 km S of El Dorado, Steyermark &
Dunsterville 104108 (NY, VEN); Depto Roscio, near Luepa, Badillo & Holmquist 6239 (MY). Delta
Amacuro: downstream from mouth of Yarakita River, Sierra Imataca, Steyermark 87446 (K, NY, SP,
VEN). Miranda: Dist. Bri?n, Rio Capaya near Sotillo, Wiersema & Gonz?lez 2205 (F, MY, NY, UNA,
VEN). Monagas: R?o Tigrito, al SSW de San Tom?, Rutkis 158 (VEN); R?o Mapirito, carretera
Matur?n-Temblador, Rutkis 159 (VEN). Zulia: Dist. Col?n, entre Boca Escalante y Pta. Pausada,
Bunting & Luis 6497 (MO).
Field studies of N. rudgeana (Cramer et al. 1975; Prance & Anderson 1976)
and observations of plants in cultivation (from Wiersema <? Gonzalez 2205) indi
cate that flowers are protogynous, opening for two or three successive nights.
Flowers reportedly open at dusk each night. The stigma is receptive, covered with
stigmatic fluid, the first night, and the stamens dehisce the second night. Flowers
both nights emit a strong "fruity" or "aniselike" odor. They are visited by the
scarab beetles Cyclocephala verticalis Burmeister and C. castanea Olivier. Al
though closure of flowers is not discussed in these studies, Conard (1905) reports
flowers to close at dawn, at 2-3 a.m., or before midnight. In view of the flowering
responses of other species of subg. Hydrocallis, it is likely that the different
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100 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
closing times given correspond to flowers of different ages. The stigma clearly
remains receptive the second night, as seeds are produced autogamously.
Nymphaea rudgeana reproduces entirely via seed production, much of this
autogamous; no form of asexual reproduction has been observed.
Based on evidence from phytochemical and crossing studies, N. amazonum
subsp. pedersenii appears to be the closest taxonomic relative to N. rudgeana
within subg. Hydrocallis. The seed morphology, general morphology, and
chromosome number of N. rudgeana all suggest a relationship to subg. Lotos and
indicate that it may be of polyphyletic origin. Nymphaea rudgeana is the only
member of subg. Hydrocallis with dentate leaves.
Plants growing in flowing water often produce an abundance of submersed
leaves. These leaves are more delicate and often have entire margins. Sagot
(1881) described one such population in French Guinea as N. amazonum var.
forma-submersa.
Nymphaea fragrans Gardner ex Caspary in Martius, Fl. bras. 4(2): 176. 1878,
pro syn.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 101
Nymphaea nocturnea March ex Hooker, Bot. Mag. 80: t.4823. 1854, pro syn.
ACKNOWLEDGMENTS
I am exceedingly grateful to Robert R. Haynes, my doctoral advisor, for his professional guid
ance, assistance, and support without which this project could not have been completed. Other
members of my dissertation committee, Robert Krai, C. Earle Smith, Jr., Joseph F. Scheiring, and
George P. Whittle, provided editorial comments on the original manuscript, as did Edward L.
Schneider for the section on seed morphology. The constructive comments of Christiane Anderson
and Daniel J. Crawford on the submitted manuscript are appreciated as well. I am also indebted to
Susan Wittlake and Jan A. Lay for preparing some of the illustrations, Steven R. Hill for providing a
Latin description and assisting with several translations, M. Carol Argo for her efforts in typing the
original manuscript, James L. Reveal for securing funds to purchase reprints, and above all Lori
Wittlake Wiersema for her emotional support throughout. I also thank those who contributed to the
research phase of this project: Harriet E. Smith-Somerville and Deborah S. Clayton for supervising
the SEM seed and pollen research, Thomas R. Bauman for aiding in the preparation of seed sections,
Marvin L. Roberts for valuable tutelage in flavonoid chemistry, Ronald G. Lindahl for answering
several karyological inquiries, Brett C. Ratcliffe for identification of Cyclocephala beetles, and fellow
graduate students Lawrence J. Davenport, Charles N. Horn, and David L. Lentz for providing a
variety of skills and services.
Field work was made possible through the generous cooperation of Angel Gonz?lez, Antonio
Krapovickas, Carmen Crist?bal, Aurelio Schinini, Ricardo Vanni, Troels Myndel Pedersen, Marcelo
de Ataide Silva, and other staff members at VEN, CTES, and IP A, as well as Richard P. Wunderlin
at USF. Appreciation is also expressed to the curators of herbaria (B, BAB, BHMH, BM, BR, C,
COL, CORD, CTES, ENCB, F, FLAS, G, GH, GUA, IAN, INPA, K, LIL, M, MBM, MO, MY,
NY, P, PH, S, SEL, SP, U, UB, UCWI, UNA, US, VEN, W, WIS) for the loan of specimens and the
staffs of several European herbaria (B, BM, G, GOET, K, M, P) for facilitating visits to examine type
material.
Financial support for field work was received from the Latin American Studies Program, Univer
sity of Alabama, and the National Science Foundation (Grant No. DEB-8111024 to the University of
Alabama). Laboratory work was aided by a grant from the National Science Foundation (No. DEB
7818402 to the University of Alabama). This is Scientific Article No. A-4331, Contribution No. 7320
of the Maryland Agricultural Experiment Station.
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102 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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APPENDIX 1
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 105
N. conardii: 2n = 28; Barinas, Venezuela, Wiersema & Gonzalez 2214 (cultivated); Chontales,
Nicaragua, Haynes 8588 (cultivated).
N. gardneriana: 2n = 28; Gu?rico, Venezuela, Wiersema 1403 (cultivated), 1412 (cultivated),
1428; Chaco, Argentina, Wiersema etal. 2247a (cultivated).
N. jamesoniana: 2n = 28; Guayas, Ecuador, Wiersema et al. 2026 (cultivated); Florida, U.S.A.,
Wiersema 2031 (cultivated); Corrientes, Argentina, Wiersema et al. 2277 (cultivated).
N. lasiophylla: 2n = 18; Alagoas, Brazil, Wiersema et al. 2293 (cultivated).
N. lingulata: 2n = 18; Maranhao, Brazil, Wiersema et al. 2308 (cultivated).
N. novogranatensis: 2n = 28; Gu?rico, Venezuela, Wiersema 1420, Wiersema & Gonz?lez 2234
(cultivated).
N. oxyp?tala: 2n = 84; Apure, Venezuela, Wiersema & Gonz?lez 2232 (cultivated).
N. prolif?ra: 2n = 18; Guayas, Ecuador, Wiersema et al. 2028; Corrientes, Argentina, Wiersema
et al. 2273 (cultivated).
N. rudgeana: 2n = 42?; Miranda Venezuela, Wiersema & Gonz?lez 2205 (cultivated).
N. tenerinervia: 2n = 20; Maranh?o, Brazil, Wiersema et al. 2307 (cultivated).
?N. amazonum x N. prolif?ra: 3n = 27; Corrientes, Argentina, Wiersema etal. 2238 (cultivated),
2249 (cultivated), Schinini & Crist?bal 14253 (this count obtained by Camilo Quarin of Corrientes,
Argentina).
APPENDIX 2
Structure and occurrence of compounds isolated from leaves of species of Nymphaea. Unless
indicated otherwise, all collection numbers are those of the author; vouchers are deposited at UNA.
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106 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
OH O
OH O OR2 O
FIG. 38. Base structures for compounds
ferols. c. Quercetins. d. Myricetins. e. Unk
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 107
Subgenus Hydrocallis.
N. amazonum subsp. amazonum. 1436 (Venezuela): C2, Ql, Q2, Ml, M5, A, B, C. 2013
(Ecuador): C2, Ql, Ml, M5, A, B, C. 2027 (Ecuador): C2, C3, Ql, Ml, M5, A, C. 2291 (Brazil): C2,
Ql, Ml, M5, A, B, C. 2221A (Venezuela): C2, Q2, Q3, A, B, C.
N. amazonum subsp. pedersenii. 2270 (Argentina): Cl, C2, C3, Ql, Q2, Q3, Q10, Qll, Ml,
M6?, M8, A, B, C.
N. amazonum subsp. amazonum x N. lasiophylla. 2296 (Brazil): C2, C3, Ql, Q2, Q4, Q7, Q8,
A, B, C, D.
N. conardii. 2214 (Venezuela): Cl, K3, K4, K5, K6. 2230 (Venezuela): 13, 14, 15, 16. 2289
(Brazil): Cl, II, 12, 13, 17,18. Haynes 8588 (Nicaragua): Cl, Kl, K2, Q9. Haynes 8612 (Nicaragua):
Cl, Kl, K2.
N. gardneriana. 1403 (Venezuela): C2, Ql, Q9, RI, R3, Ml, M2. 1412 (Venezuela): Ql, Q9,
Qll, Ml. 1428 (Venezuela): Ql, Q2, Q9, R2, R3, mQ3. 2210 (Venezuela): Ql, Q9, R2, Ml. 2247A
(Argentina): C2, Ql, Q9, R2.
N. jamesoniana. 1343 (Mexico): Ql, Q9. 2030 (Ecuador): Ql, Q9. 2032 (Florida): Ql, Q9. 2265
(Argentina): Ql, Q9. 2290 (Brazil): Ql, Q9.
N. lasiophylla. 2293 (Brazil): M2, M7, mM3, mM4, A, B, C, FI, F2, F3, F4, mQ4. 2316 (Brazil):
M2, M7, mM3, mM4, A, B, C, FI, F2, F3, F4, mQ4.
N. lingulata. 2283A (Brazil): C3, mQl, mQ2, mMl, mM2, mM3, mM5, A, B, C. 2308 (Brazil):
mQl, mQ2, mMl, mM2, mM3, mM5, A, B, C.
N. novogranatensis. 1404 (Venezuela): C2, Ql, Q5, Q6, Q10, Qll, A, B, C. 1411 (Venezuela):
C2, Ql, Q5, Q6, Q9, Q10, Qll, A, B, C. 1420 (Venezuela): C2, Ql, Q5, Q6, Qll, A, B, C. 2234
(Venezuela): C2, Ql, Q5, Q6, Q9, Qll, A, B, C.
N. oxypetala. 2232 (Venezuela): Ql, Q9, mQ2?, Ml, A, B, C.
N. prolif?ra. 2028 (Ecuador): C2, Ql, Q8?, Qll, Ml, M4, M8, A, B, C. 2241 (Argentina): C2,
Ql, Q8?, Qll, Ml, M4, M8, A, B, C. 2273 (Argentina): C2, Ql, Q8?, Qll, Ml, M4, M8, A, B, C.
N. prolif?ra x N. amazonum subsp. pedersenii? 2238 (Argentina): C2, Ql, Qll, Ml, M4, M8,
A, B, C. 2240 (Argentina): C2, Ql, Q10, Qll, Ml, M4, M6, M8, A, B, C. 2249 (Argentina): C2, Ql,
Qll, Ml, M4, M8, A, B, C. 2254 (Argentina): C2, Ql, Qll, Ml, M4, M8, A, B, C. 2255 (Argen
tina): C2, Ql, Qll, Ml, M4, M8, A, B, C.
N. rudgeana. 2205 (Venezuela): Cl, C2, C3, Q2, Q3, Q9, A, B, C. 2285 (Brazil): Cl, C2, C3,
Ql, Q2, Q3, Q9, A, B, C. 2299 (Brazil): Cl, C2, Ql, Q2, Q3, Q9, A, B, C.
N. tenerinervia. 2307 (Brazil): C2, Ql, Ml, M5, M6, A, C. 2310 (Brazil): C2, Ql, Q10, Ml, M6,
A, B, C. 2311 (Brazil): C2, Ql, Q10, Ml, M5, M6, A, B, C. 2314 (Brazil): C2, Ql, Ml, M5, M6, A,
B, C.
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108 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
Subgenus Brachyceras.
N. ampia. 1345 (Mexico): C2, Ql, Q9, Q10. 1452 (Florida): C2, Ql, Q9, Q10, mQ2, mMl,
mM3, A, C. 2021 (Ecuador): C2, Ql, Q9, Q10, mQ2, A, C.
N. elegans. 1981 (Florida): C2, Q9, Q10, mQ2, M3, M6, mMl, mM3, A, B.
Subgenus Lotos.
N. lotos. 1442 (Florida): Cl, C2, M6, A, B, C.
Subgenus Nymphaea.
N. candida. 2196A (Germany): Q9, mQ2, M6, mM3, A, B.
N. mexicana. 281 (Alabama): C2, Ql, Qll.
N. odorata. 274 (Alabama): C2, Q9, mQ2, Ml?, M3, M6, mMl, mM3, A, B.
The numbers in parentheses refer to the corresponding species in the text and in the Numerical
List of Species presented above.
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1987 NYMPHAEA SUBGENUS HYDROCALLIS 109
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110 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 16
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NYMPHAEA SUBGENUS HYDROCALLIS 111
2270 (10b); 2273 (11); 2277 (1); 2284 (10a); 2310 (9); 2311 (9); 2312 (10a); 2313 (12);
2285 (14); 2286 (14); 2287 (10a); 2289 (2); 2314 (9); 2315 (1); 2316 (12); 2317 (12).
2290 (1); 2291 (10a); 2293 (12); 2294 (12); Woodson et al. 1225 (2).
2295 (12); 2297 (1); 2298 (12); 2299 (14); Woolston, A. L. 1257 (3).
2300 (1); 2302 (12); 2303 (10a); 2304 (10a); Wright, C. 2491 (10a); 3192 (2).
2306 (1); 2307 (9); 2308 (13); 2309 (14); Wright, S. 1 (12); 4 (12).
Accepted names are in roman type; the main entry for each is in boldface. Synonyms are in
italics.
Aesculus Linnaeus 44 12, 15, 21, 22, 23, 24, 26, 27, 28, 31, 43,
Barclaya Wallich 12 45, 46, 96, 108
Brasenia Schreber 24 subg. Hydrocallis (Planch?n) Conard 49-50
Cannabis Linnaeus 44 subg. Lotos (de Candolle) Conard 3, 5, 7, 8,
Castalia Salisbury 2, 48 12, 15, 21, 23, 24, 25, 26, 27, 28, 29, 32,
amazonum (Martius & Zuccarini) Britton & 37, 43, 44, 45, 46, 47, 48, 100, 108
P. Wilson 80 subg. Nymphaea 3, 4, 8, 9, 11, 12, 15, 21, 22,
ampia Salisbury 49, 85 23, 25, 26, 27, 28, 31, 43, 44, 45, 46, 47,
blanda (G. Meyer) Lawson 62 96, 108
gibertii Morong 51 subsect. Hydrocallis (Planch?n) Caspary 49
jamesoniana (Planch?n) Britton & P. Wilson tribe Chamaehtos Lehmann 49
51 alba Linnaeus 2, 25, 37, 44, 48, 65
rudgeana (G. Meyer) Britton & P. Wilson 96 albo-viridis Saint-Hilaire 81
Cyclocephala Latreille 32, 35 amazonum Martius & Zuccarini 2, 3, 8, 9, 12,
castanea Olivier 32, 65, 99 15, 25, 29, 32, 33, 34, 36, 38, 39, 40, 41,
mollis Endr?di 32, 86 42, 43, 44, 48, 49, 51, 65, 80-87, 90, 101
p?trida Burmeister 35 f. goudotiana (Planch?n) Caspary 82
verticalis Burmeister 32, 35, 57, 85, 99 subsp. amazonum 8, 17, 27, 33, 35, 81-85,
Euphorbia Linnaeus 44 86, 87, 93, 104, 107, 108
Euryale Salisbury 37 subsp. pedersenii Wiersema 8, 9, 18, 30,
ferox Salisbury 37 31, 33, 43, 44, 48, 81, 83, 84, 86-87, 90,
Hydrostemma Wallich 12 100, 104, 107, 108
Leuconymphaea Ludwig ex Kuntze 48 var. forma-submersa Sagot 85, 96, 100
amazonum (Martius & Zuccarini) Kuntze 80 var. goudotiana (Planch?n) Conard 82
blanda (G. Meyer) Kuntze 62 ampia (Salisbury) de Candolle 2, 12, 15, 22,
gardneriana (Planch?n) Kuntze 52 23, 26, 28, 39, 40, 43, 44, 49, 51, 59, 85,
goudotiana (Planch?n) Kuntze 82 100, 101, 108
jamesoniana (Planch?n) Kuntze 51 var. rudgeana (G. Meyer) de Candolle 96
lasiophylla (Martius & Zuccarini) Kuntze 90 var. speciosa (Martius & Zuccarini) Cas
oxyp?tala (Planch?n) Kuntze 71 pary 100
Nelumbo Adanson 2 belophylla Trickett 3, 7, 8, 24, 31, 43, 47, 48,
Nuphar Smith 2 50, 61, 65-68, 71, 73, 108
lutea (Linnaeus) Smith 37 blanda G. Meyer 2, 3, 12, 62, 63, 65
Nymphaea Linnaeus 48-49 var. amazonum (Martius & Zuccarini)
sect. Castalia (Salisbury) de Candolle 2 Planch?n 80
sect. Hydrocallis Planch?n 2, 49 var. fenzliana (Lehmann) Caspary 62
sect. Lotos de Candolle 2 caerulea Savigny 37
sect. Lytopleura Caspary 2 candida Presl 25, 37, 39, 40, 43, 44, 108
sect. Symphytopleura Caspary 2 candolleana Lehmann 82, 85
subg. Anecphya (Caspary) Conard 12, 21, 24, capensis Thunberg 26, 28, 37, 100
25, 26, 27, 31, 45, 46 conardii Wiersema 7, 8, 9, 11, 12, 15, 18, 19,
subg. Brachyceras (Caspary) Conard 3, 5, 9, 21, 28, 29, 30, 32, 34, 35, 36, 38, 39, 40,
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112 SYSTEMATIC BOTANY MONOGRAPHS
41, 42, 47, 51, 55-58, 61, 65, 66, 77, odorata Ait?n 4, 15, 22, 23, 25, 26, 28, 31,
104, 107, 108 39, 40, 43, 44, 80, 100, 108
dentata Schumacher 25 oxypetala Planch?n 3, 7, 8, 9, 10, 12, 18, 20,
elegans Hooker 15, 22, 23, 26, 28, 31, 39, 40, 27, 30, 31, 35, 38, 39, 40, 43, 46, 47, 50,
43, 44, 108 65, 68, 69, 71-74, 104, 107, 108
fenzliana Lehmann 62, 65 passiflora Lehmann 62
foetida Gardner ex Lehmann 100 potamophila Wiersema 7, 8, 10, 18, 20, 31,
fragrans Gardner ex Caspary 100 43, 47, 50, 65, 68-71, 72, 108
gardneriana Planch?n 3, 7, 8, 9, 17, 18, 28, prolif?ra Wiersema 8, 9, 11, 18, 24, 27, 29,
29, 30, 34, 35, 36, 38, 39, 40, 41, 42, 47, 30, 31, 32, 33, 34, 36, 38, 39, 40, 41, 42,
51, 54, 58-62, 65, 66, 96, 104, 107, 108 48, 51, 76, 81, 87-90, 91, 93, 104, 107,
gibertii (Morong) Conard 2, 51 108
gigantea Hooker 25, 31 raja Lehmann 71, 73
glandulifera Rodschied 7, 8, 9, 12, 18, 20, 21, rubra Roxburgh 37
27, 31, 32, 34, 43, 47, 51, 58, 62-65, 69, rudgeana G. Meyer 2, 3, 6, 7, 8, 9, 10, 12,
72, 96, 108 13, 15, 16, 23, 26, 27, 29, 31, 32, 34, 36,
goudotiana Planch?n 81, 85 37, 38, 39, 40, 43, 46, 48, 50, 62, 81, 85,
integrifolia Salzmann ex Lehmann 100 96-100, 104, 107, 108
jamesoniana Planch?n 3, 7, 8, 9, 11, 12, 13, var. amazonum (Martius & Zuccarini)
18, 19, 21, 23, 26, 28, 29, 30, 32, 34, 35, Grisebach 80
36, 37, 38, 39, 40, 42, 47, 50, 51-55, 58, sagittariaefolia Lehmann 51, 55
61, 63, 65, 77, 104, 107, 108 sinuata Salzmann ex Lehmann 96, 101
lasiophylla Martius & Zuccarini 2, 3, 7, 8, 9, stellata Willdenow 25
11, 15, 16, 26, 27, 29, 30, 35, 38, 39, 40, stenaspidota Caspary 3, 59
42, 43, 44, 45, 48, 50, 80, 81, 85, 90-94, tenerinervia Caspary 3, 8, 9, 12, 15, 17, 27,
96, 104, 107, 108 29, 30, 32, 33, 34, 35, 36, 38, 39, 40, 42,
leiboldiana Lehmann 100 43, 48, 51, 59, 65, 76, 77-80, 81, 90, 94,
lingulata Wiersema 7, 8, 9, 15, 16, 26, 27, 29, 100, 104, 107, 108
30, 32, 33, 34, 35, 36, 37, 38, 39, 40, 42, tetragona Georgi 25
43, 44, 48, 50, 93, 94-96, 104, 107, 108 tropaeolifolia Lehmann 101
lotus Linnaeus 4, 15, 23, 24, 25, 26, 28, 36, viol?cea Lehmann 25
37, 39, 40, 43, 44, 108 wittiana Ule 3, 59
maximiliani Lehmann 80, 100 Oenothera Linnaeus 44
mexicana Zuccarini 12, 25, 26, 28, 31, 39, 40, Ondinea Hartog 12, 24
46, 108 Victoria Lindley 37
nocturnea March ex Hooker 101 amaz?nica (Poeppig) Sowerby 37
novogranatensis Wiersema 8, 9, 14, 18, 19, regia Lindley 37
21, 27, 29, 30, 32, 33, 34, 35, 36, 37, 38,
39, 40, 42, 43, 47, 48, 51, 74-77, 78,
104, 107
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