Ovary, Fruit, and Seed Morphology of the Lythraceae

Author(s): Shirley A. Graham and Alan Graham

Source: International Journal of Plant Sciences, Vol. 175, No. 2 (February 2014), pp. 202-240
Published by: The University of Chicago Press
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Int. J. Plant Sci. 175(2):202–240. 2014.
䉷 2014 by The University of Chicago. All rights reserved.
1058-5893/2014/17502-0005$15.00 DOI: 10.1086/674316

OVARY, FRUIT, AND SEED MORPHOLOGY OF THE LYTHRACEAE

Shirley A. Graham1,* and Alan Graham*

*Missouri Botanical Garden, 4344 Shaw Boulevard, Saint Louis, Missouri 63110, USA

Editor: Larry Hufford

Premise of research. The ovary, fruits, and seeds of the Lythraceae present an array of morphological
characters that have not been fully explored for their utility in clarifying taxonomic, phylogenetic, and historical
biogeographic questions. This study provides a morphological record of ovary, fruit, and seed characters for
the family that can be used to address these questions. The seeds, in particular, display diverse adaptations
to dispersal and are emphasized in the study. A more complete knowledge of the fruits and seeds of the
Lythraceae further promises to aid identification of lythracean fossil fruit and seed remains from late Cretaceous
and Cenozoic floras around the world.
Methodology. The morphology of alcohol-preserved and softened dried ovaries, fruits, and seeds is de-
scribed based on hand sections studied using LM. SEM further details the external features of mature seeds
of all genera of the family. Selected character states are shown on a previously produced molecular-based
phylogeny to examine distribution of adaptations that influenced the radiation and diversification of the family.
Pivotal results. Diversity in the mostly superior ovary of the family includes forms with elongated or
globose axile placentas, ovarian stipes, and differing degrees of septal development. Tropical tree genera show
tendencies toward an inferior ovary; semi-inferior to fully inferior ovaries occur in Punica and Trapa. The
most common fruit type, occurring in 10 of the 28 genera, is a dry, thin- to thick-walled capsule of two to
five locules that splits irregularly to free the seed. Other capsules open loculicidally, septicidally, by a circum-
scissile opening, or are indehiscent. Capuronia, Punica, and Sonneratia produce dissimilar indehiscent berrylike
fruits; a drupe is unique to Trapa. Seed size in the Lythraceae ranges widely, with 18 of 28 genera producing
seeds less than 1 mm long. The largest seeds are winged seeds in Lagerstroemia (to 1.6 cm) and Lafoensia
(to 3.5 cm). Seed shape is obpyramidal to bilateral, and seed coats, composed mainly of polygonal or rectangular
cells, are smooth surfaced with a reticulate or scalariform pattern. Aerenchymatous float tissue, wings, and
unique inverted mucilaginous trichomes in the seed coat that evaginate upon wetting to form long “hairs”
are specialized features of adaptive significance for dispersal, germination, and establishment.
Conclusions. Ovaries, fruits, and especially the seeds are morphologically well diversified in the Lythraceae.
Character states are phylogenetically autapomorphic on terminal branches or homoplastic and do not support
the major lineages or clades. The relatively extensive fossil record of the family, including seeds of Decodon
(73.5 Ma) and pollen of Lythrum/Peplis (82–81 Ma), suggests early adaptations of seeds for aquatic dispersal
by development of extensive float tissue (Decodon) and for enhanced dispersal and germination by inverted
mucilaginous seed coat trichomes (inferred from extant Lythrum/Peplis). The adaptations are responsible in
large part for the present global distribution of the family.

Keywords: dispersal, fruits, Lythraceae, phylogeny, seed morphology, trichomes.

Introduction acters on molecular phylogenies has provided new insight into

Comparative morphology of reproductive organs has long
been central to the study of relationships among and within
angiosperm families (Kaplan 2001). The seed surface, espe-
cially, provides characters of taxonomic significance (Boese-
winkel and Bouman 1984, 1995), attested to by the multitude
of studies employing seed characters in classification (Jensen
1998). In recent years, the optimization of morphological char-
1
Author for correspondence; e-mail: shirley.graham@mobot.org.
Manuscript received March 2013; revised manuscript received August 2013;
electronically published January 10, 2014.
patterns of evolutionary change in plants. Still, comparative
morphology of ovaries, fruits, and seeds in many plant families
is superficial at the generic level even while multigene molec-
ular relationships of genera and species are increasingly
known, and this is the case with the Lythraceae.
The Lythraceae are a family of nearly cosmopolitan distri-
bution, with greatest presence in the subtropics and tropics of
both hemispheres. The 28 genera (Graham et al. 2011) exhibit
an array of ovary, fruit, and seed characters not yet fully ex-
plored for their utility in clarifying taxonomic, phylogenetic,
and historical biogeographic questions. In particular, detailed
descriptions and illustrations of the very small seeds common

202

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY
to the majority of genera have not been made in spite of their
apparent diversity. A goal of this study is to make available a
morphological record of ovary, fruit, and seed character states
for the family that can be utilized for systematic and phylo-
genetic purposes and for the identification of fossil remains.
The Lythraceae have an extensive fossil record that includes
flowers, fruits, and very small seeds—many less than 2 mm
long—that are easily overlooked and difficult to identify given
lack of adequate descriptions (Graham 2013). New informa-
tion presented here will aid recognition of both modern and
fossil Lythraceae fruits and seeds and allow increased under-
standing of the evolution and global history of the family.
Koehne (1884, 1885, 1903) in his monographic studies of
the Lythraceae described vegetative morphology (Koehne
1884) and floral structure (for the gynoecium, see Koehne
1885, pp. 30–34) for the 22 genera known at that time and
provided references to earlier nineteenth-century descriptions.
More recent observations are included in floral studies by Tobe
et al. (1998) and others (e.g., Cuphea calophylla Cham. &
Schltdl. [Schoenberg and Hofmeister 1986; Schoenberg 1989];
Trapa natans [Titova et al. 1997]; Cuphea carthagenensis
(Jacq.) Macbr. [Lusa and Bona 2011]; Ammannia, Nesaea, and
Hionanthera [Graham et al. 2011]).
Early descriptions were primarily concerned with unusual
“hairs” that appeared on some seed surfaces upon wetting (as
in Lythrum thesioides, Lythrum portula [Koehne 1878]; Cu-
phea viscosissima [Klebs 1885]; Cuphea lanceolata, Lythrum
salicaria [Brandza 1889, 1891]; C. viscosissima [Correns
1892]; five species of Cuphea, L. salicaria, and Heimia [Gruet-
ter 1893]). Generic-level surveys of angiosperm seeds include
some Lythraceae (Netolitzky 1926; Corner 1976; Takhtajan
1996; Kirkbride et al. 2006), and revisions and recently de-
scribed genera provide fruit and seed information (Ammannia
[Graham 1985]; Rotala L. and Ammannia L. [Panigrahi 1986];
Decodon, Heimia, and Nesaea [Graham 1977]; Ginoria [Gra-
ham 2010]; Koehneria [Graham et al. 1986]; Lafoensia [Lour-
teig 1985]; Lourtella [Graham et al. 1987]; Tetrataxis [Graham
and Lorence 1978]; Woodfordia [Graham 1995]). Morpho-
logical features of the ovaries, fruits, and seeds of the majority
of the genera, however, have not been described or compared
across the family.
The classification of the Lythraceae by Koehne (1903) is still
the classification in use, and it relies heavily on ovary, placen-
tal, and seed characters. Koehne recognized two tribes, Lyth-
reae Koehne and Nesaeeae Koehne. In tribe Nesaeeae, septa
were considered complete, and the placenta always continuous
with the style; in Lythreae, septa were “interrupted,” and the
placenta was not continuous with the style. All septa in the
family are now known to be incomplete, i.e., not completely
fused to each other or to the apex of the ovary. Koehne further
divided tribe Lythreae into two subtribes based on seeds emar-
ginate (or if marginate, then flowers zygomorphic) versus seeds
dorsally compressed and encircled by a wing. He similarly
divided tribe Nesaeeae into two subtribes based on seed coats
not thickened or winged versus seed coats winged or with a
spongy apex. In most genera, seeds were described only as
“numerous” or “very numerous” and “minimal” or “small”
(Koehne 1903). Neither the tribes based on ovary morphology
nor the subtribes based on seed morphology are supported by
molecular evidence (Graham et al. 2005).
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203
The role accorded to these features in the classification of
the family has not been examined critically or described at
more than a superficial level. Thus, the diversity of some major
floral organs has yet to be fully utilized in reconstructing the
evolutionary history of the Lythraceae or revising the now
outdated taxonomic classification. In this study, we report re-
sults of a morphological survey of the ovary, fruit, and seeds
of all lythraceous genera to provide a more complete char-
acterization of the family and for use in future phylogenetic,
taxonomic, and paleobotanical/historical biogeographic
research.
Methodology
Descriptions are based primarily on herbarium and liquid-
preserved material of flowers, fruits, and seeds and greenhouse
collections at the Missouri Botanical Garden (MO), Saint
Louis. Multiple collections of each genus were used to limit
removal of material from any one specimen and to ensure that
descriptions included any significant morphological variations.
The number of collections available for each genus differed,
as did the state of the material available. For the largest genera,
the descriptions are considered representative but not inclusive
of all variants at the species level. Voucher information is pro-
vided for most illustrated material; vouchers are deposited at
MO unless otherwise noted. Herbarium material was softened
in water with a drop of detergent and warmed in a microwave
for 10 s or less, and observations of most ovaries and fruits
were made from hand dissections under a dissecting micro-
scope; some permanent anatomical sections in the authors’
collections and photographs of sections provided by H. Tobe,
University of Kyoto, Japan, were also employed to assist in
the morphological interpretations. Seed descriptions were doc-
umented by LM and SEM. Seeds were sputter-coated with
gold-palladium for 120 s and examined using Cambridge Ste-
reoscan Mark IIA and S420 scopes or a Nikon/JEOL Neoscope
JCM-5000. Seed length was measured as the longest distance
from the proximal end to the distal end of the seed including
wings if present; seed width was measured as the greatest dis-
tance from outer margin to outer margin of the seed measured
at right angles to the proximal/distal dimension. Anticlinal cell
walls are those perpendicular to the seed surface, and periclinal
cell walls are parallel to the seed surface.
For purposes of phylogenetic discussion, a previously gen-
erated majority-rule Bayesian consensus of 9000 trees was uti-
lized with slight modification (Morris 2007). The tree is based
on combined sequence data from the rDNA nuclear internal
transcribed spacer (ITS) and chloroplast rbcL, trnL-F spacer,
trnK-matK region, and atpB-rbcl intergenic spacer (Morris
2007). Three major lineages and six clades are recognized. The
phylogeny carries a high degree of confidence with well-sup-
ported branches in all but the basal trichotomy of the tree
where the relationship of the three major lineages remains
unresolved. The tree is modified for this study by collapse of
the Heimia-Rotala-Didiplis clade to the base of the tree because
of the low support values for that branch (posterior probability
55; maximum likelihood support 60%). No molecular data
were available for Crenea; the other 27 genera of the family
are represented. The positions of Cuphea (Graham et al. 2005;
Barber et al. 2010) and Diplusodon (T. Cavalcanti and P. Inglis,
 Fig. 1 Ovaries, placentas, fruits. A, Lafoensia pacari A.St.-Hil., longitudinal section of a young flower, stipitate superior ovary surrounded
by floral tube on which are inserted numerous stamens with long filaments. 1, Floral cup; 2, ovary; 3, ovary stipe. B, Adenaria floribunda,
longitudinal section of young fruit. 1, Black resin-filled trichomes visible on capsule surface; 2, globose fertile portion of placental axis,

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY 205

unpublished data) are added based on a tree that found the
same lineages and clades as the Morris tree. The position of
Decodon and Lythrum in all analyses is unstable, occurring
either as the earliest-diverging branch within the Lythraceae
or, as in the Morris tree, on a moderately supported branch
(Bayesian prior probability 78; maximum likelihood support
74%) as sister to the major clade Duabanga-Ammannia. Se-
lected character states were parsimoniously optimized on the
previously published phylogeny employing MacClade (Mad-
dison and Maddison 1992) with characters unordered and
equally weighted under Acctran transformation. The outgroup
consisted of Ludwigia, Fuchsia (Onagraceae, sister to Lyth-
raceae), and Combretum (Combretaceae) based on family-level
phylogenetic relationships in the Myrtales (Sytsma et al. 2004).
Commonly Held Features in Lythraceae Ovary,
Fruit, and Seed Morphology
The ovaries, fruits, and seeds of the Lythraceae develop, with
few exceptions, in actinomorphic, hermaphroditic, perigynous,
four- to six-merous, diplostemonous flowers. The floral cup
or floral tube (hypanthium) is persistent. The gynoecium con-
sists of a superior, ellipsoid, subglobose, or globose two- to
multicarpellate ovary, single style, and capitate to punctiform
stigma. The ovary is smooth surfaced, rarely bearing trichomes
around the apex, and it can be depressed apically or appear
fluted in transverse section, the flutes equaling the number of
locules. In some genera the ovary is clearly stipitate (fig. 1A).
Semi-inferior to inferior ovaries are present in a few genera.
The number of locules varies mostly from two to five or six;
three genera previously recognized in the satellite families Dua-
bangaceae, Punicaceae, and Sonneratiaceae have more. There
is a trend toward zygomorphy that is most fully expressed in
Cuphea P. Browne, the largest genus of the family with ∼250
species.
The placenta is a solid, central axis except in Duabanga and
Sonneratia, where it is hollow (Tobe et al. 1998). Placentation
is basically axile; however, in many genera it appears free-basal
at fruit maturity as a result of breakage of the placental con-
nection above the ovule-bearing placental lobes and reduction
or cessation of the septa above the ovule-bearing lobes (fig.
2). The placenta may remain elongate and slender as the ovary
enlarges, with ovules directly attached to the central axis, or
lobes of ovule-bearing tissue may protrude into the locules,
enlarging to form a subglobose to globose cushion covered
with many minute to small seeds (fig. 1B, 1C). The basal por-
tion of the axis can be sterile forming a placental stipe (fig.
1D). Septa cease attachment to the placenta at various levels
from the base of the ovary to the ovary apex depending on
the genus.
Fruits are primarily capsules. Fruit walls range from thin,
dry, and fragile (fig. 1D) to leathery and often ultimately in-
durate (fig. 1E). Thin walls are defined here as five cell layers
thick or less; thick walls are more than 10 cell layers; mod-
erately thick walls are six to 10 cell layers thick. These are
estimates based on observations of hand sections under LM;
the number of cell layers varies in different parts of the fruit,
and anatomical thin sections are required to more precisely
determine the full variation.
The seeds of the Lythraceae develop from anatropous, bi-
tegmic ovules in which the ovules nod on a funicle. In the
Lythraceae the funicle is usually short, resulting in close align-
ment of the hilum and micropyle at the funicular end of the
seeds (fig. 3). The seeds in this study are illustrated with the
funicle and micropyle oriented at the proximal end and the
chalaza at the distal end of the seed. Seed shape and seed
outline are also described based on this orientation. With re-
spect to the attachment of the seed to the placenta, seeds project
outward in all directions from a globose placenta (fig. 1B, 1C)
or are arranged in two irregular rows in each locule (fig. 1D,
1E). In the past, confusion has arisen in describing some seed
orientations in relation to the placenta (for Decodon and fossil
relatives, see Tiffney 1981). The orientation of the seed has
been variously described as adaxial/abaxial, dorsal/ventral, or
raphal/antiraphal. The orientation is described here as raphal/
antiraphal, and the raphal side of the seed faces the placental
axis.
The majority of genera produce seeds classified as small to
medium in size, 3 mm long or less (Werker 1997). The embryo
fully occupies the interior of the seed; no endosperm is present
at seed maturity. The embryo is erect, with the apex of a short
radicle directed toward the proximal end of the seed. The
cotyledons, thickened by oil and protein storage products, in-
vest the radicle for at least half its length (Graham and Kleiman
1987). Trapa is unique in the family in producing a single
starchy seed per fruit.
Morphological diversity of the seeds is found primarily in
size, shape, and presence of adaptations for enhanced disper-
sal. Uniquely in the angiosperms, some genera of the Lyth-
raceae form internal trichomes in the epidermal cells of the
seed coat. These swell upon becoming wet and evaginate from
the cell to cover the seed with a “hairy” coat of mucilaginous

discontinuous with the ovary apex, placentation ultimately free-basal, basal portion of placenta sterile, stipe-like, not clearly visible in photo;
3, persistent floral cup subtending the capsule. C, Capuronia benoistii, longitudinal section, young stage of berrylike fruit, prior to presence of
extensive mucilage. 1, Seeds attached on expanded lobe of the placenta, upper portion of the placenta still continuous with the ovary apex; 2,
one partially mature seed and four undeveloped seeds visible attached to placental lobe; 3, thick wall and complete septum in locule on the
right; 4, remnants of floral cup. D, Cuphea llavea Lex., longitudinal section of floral tube and enclosed thin-walled capsule. 1, Persistent floral
tube; 2, thin capsule wall; 3, threadlike remnant of one of the two vestigial septa (at tip of pointer); 4, opened capsule wall, extended to the
right; 5, elongate placenta with proximal sterile stipe, discontinuous distally above seed-bearing portion, some seeds removed from the dorsal
locule to expose the placenta, base of funicles of removed seeds visible. E, Lagerstroemia indica, longitudinal section through indurated capsule.
1, Seed wing directed upward, wings of two lower seeds on left obscured by wing of upper seed; 2, attachment point of seed to the placenta
(at tip of pointer); 3, persistent floral cup; 4, complete septum extending to apex of ovary, septa not fused together at ovary apex; 5, elongate
placenta, extending to ovary apex. Scale bar p 0.5 mm in A, 1 mm in B–D, 2 mm in E. (A from Cavalcanti et al. 2326; B from Wiley 504;
C from Chavet 261; D from Tenorio 10175; E from cultivated material.)

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 Fig. 2 Sectioned young flowers of Rotala, Ammannia, Didiplis, and Pleurophora prior to seed maturity, illustrating diversity in ovaries,
placenta, and, in Rotala, incomplete septal partitioning. A–C, Rotala rotundifolia (Roxb.) Koehne. A, Longitudinal section (LS); elongate structures
at far left and right are bracteoles; internal to bracteoles the floral cup encloses the subglobose ovary; base of the ovary is flanked by dark-
stained nectaries; the globose placenta is stipitate, and seeds cover the surface; placentation appears free-basal at maturity. B, Transverse section
(TS) at approximately midlevel of ovary; at this level septa extend from ovary wall to central placental axis; four stamen filaments visible outside
the septa; the floral cup and a pair of bracteoles are to the outside of the ovary. C, TS just above apex of globose placenta; at this more distal
level in the ovary the septa are incomplete. D, Ammannia coccinea Rottb. LS; sessile ovary enclosed in a floral cup; placenta ellipsoid, short-
stipitate, distal connection to ovary apex broken. E, Didiplis diandra. LS; ovary sessile; dark-stained nectaries visible at base of floral tube;
placenta globose; placentation free-basal at maturity. F, Pleurophora saccocarpa Koehne. LS; young flower with placental connection to ovary
apex still intact, bilocular ovary included in an elongate floral tube, basal remnants of floral tube present to left and right of ovary; ovary stipe
encircled by the nectary; zygomorphy expressed by unequal locules and placental lobes. Anatomical sections prepared and photographed by H.
Tobe, used with permission (see additional genera in Tobe et al. 1998).

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY
Fig. 3 Generalized diagram of an anatropous ovule in relation to
seed morphology of a lythraceous seed. A, Generalized ovule structure.
B, Bilateral seed of Cuphea hookeriana Walp. 1, Distal end of raphe
in the funicle at the chalazal (i.e., terminal) end of seed. 2, Chalazal
area, not visible on seed surface. 3, Micropyle formed by outer and
inner integuments, obscured on the seed at proximal seed margin by
surrounding tissues. 4, In A, the funicle, occurring in close association
with the micropylar opening at the proximal end of the seed; in B,
hilum, the scar produced following detachment of the mature seed
from the funicle.
threads (e.g., figs. 4E, 15D). Seeds of the Lythraceae possess
a characteristic internal hardened pellicle. Anatomically, this
consists of thick-walled inner sclerotic layers developed from
the outer integument and an adjoining lignified cell layer of
the inner integument. One or both of these specialized layers
also occur in the Onagraceae, sister family to the Lythraceae,
and in the Combretaceae, sister to the rest of the order Myr-
tales (Corner 1976; Tobe et al. 1987; Sytsma et al. 2004).
Generic Descriptions
Adenaria Kunth
One species, Adenaria floribunda Kunth; small trees or tall
shrubs, flowers actinomorphic, weakly trimorphic (Nevling
1958) and incipiently dioecious; southern Mexico to Argen-
tina; evergreen forests, disturbed secondary forests. Figures 1B,
4A–4D.
Ovary (of fertile female flowers). Superior, stipitate with
stipe to ∼1.0 mm, ovary obovoid to orbicular, 2 (3) locular;
placental stipe present, 1–1.4 mm, placenta globose, lobes pro-
truding into the locules; septa thin, ceasing at ovary apex;
placenta free-basal at maturity; ovules ∼70–80.
Fruits. Capsules, 3–7 mm # 3–4 mm, globose, light
brown to brown, externally bearing sessile, globose, black
resin-filled trichomes, sometimes also pubescent on distal half
of capsule; pericarp moderately thick, indurated, brittle, fruit
well exserted from the persistent floral cup at maturity, irreg-
ularly splitting.
Seeds. Approximately 7–20 per capsule, 0.9–1.3 mm #
0.8–1.2 mm, golden brown, covering globose surface of pla-
centa, tightly packed; seed shape obpyramidal when immature,
becoming subglobose, slightly bilateral, 3–5 (6) faceted, raphal
face 2 faceted or slightly flattened, raphe inconspicuous on
seed surface, antiraphal face typically convex, 3–4 faceted or
rounded; seed outline obovate to suborbicular, apex rounded
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207
or obtuse; seed surface smooth; raphal face irregularly retic-
ulate, cells irregularly rectangular to polygonal, anticlinal walls
prominent, periclinal walls depressed; antiraphal faces irreg-
ularly reticulate, cells smaller and more polygonal/hexagonal
than on raphal face, anticlinal walls less prominent, periclinal
walls smooth, slightly depressed.
Epidermal cell internal trichomes. Absent.
Ammannia L.
Approximately 80–100 species (including Nesaea Comm. ex
Kunth and Hionanthera A. Fern. & Diniz; Graham and Gan-
dhi 2013); annual or perennial herbs, terrestrial, amphibious,
or aquatic, flowers actinomorphic, hermaphroditic, some di-
or tristylous; primarily Africa and Asia, seven species distrib-
uted in tropical to temperate America; fresh or brackish
marshes, margins of ponds and rivers. Figures 2D, 4E–4H,
18A.
Ovary. Superior, sessile, nonstipitate, globose to oblong,
(1) 2–4 (5) locular; placenta nonstipitate or stipe less than 0.5
mm, globose to narrowly ellipsoid, infrequently slender-elon-
gate, smooth or 2–4 sulcate, slightly fluted in cross section,
lobes protruding into the locules; septa thin, ceasing near ovary
apex in species with elongate placenta, ceasing just above
ovule-bearing portion in species with globose or oblong pla-
centa, placenta typically free-basal at maturity; ovules typically
100⫹.
Fruits. Capsules, 1–5 mm # 1–6 mm, globose to oblong,
golden brown to red, glabrous; pericarp thin, transparent;
fruits included or partially exserted from persistent floral tube
at maturity, irregularly splitting from apex or initially circum-
scissile and then irregularly splitting.
Seeds. One hundred or more, often 250–300⫹ per cap-
sule, 0.3–0.4 mm # 0.3–0.5 mm, golden brown to brown or
dark red, densely covering surface of the globose placenta (ex-
cept in Ammannia mossambicensis [A. Fern. & Diniz]
S.A.Graham & Gandhi only 2–5 seeds per capsule, 1.0–1.8
mm # 0.7–1.3 mm, dark violet); seed shape obovoid-sub-
globose, raphal face bearing aerenchymatous float tissue, con-
cave after collapse of float tissue (float absent on A. mossam-
bicensis), raphe inconspicuous on seed surface, antiraphal face
convex; seed outline obovate, elliptical, oblong, or suborbi-
cular, apex rounded; seed surface smooth, cells of float tissue
thin walled, reticulate to scalariform, rectangular, anticlinal
walls straight, slightly raised, periclinal walls smooth, flat to
slightly depressed, cells of antiraphal face reticulate, rectan-
gular, anticlinal walls straight or sinuous, periclinal walls flat
or slightly depressed.
Epidermal cell internal trichomes. Present, ∼70–110 mm
(294 mm in A. mossambicensis), straight, deeply or shallowly
ribbed, lightly to densely tuberculed, tubercles circular or
elongate.
Capuronia Lourt.
One species, Capuronia benoistii (Leandri) P.E.Berry; spiny
shrubs, flowers actinomorphic, incipiently dioecious, func-
tionally unisexual, anther-bearing flowers preserving a rudi-
mentary ovary; endemic to Madagascar; dry deciduous forests.
Figures 1C, 5A–5C.
Ovary (of fertile flowers). Superior, sessile, nonstipitate,
 Fig. 4 A–D, Adenaria floribunda. A, Antiraphal face; scale bar p 0.5 mm. B, Raphal face; scale bar p 0.5 mm. C, Oblique view, raphal
face to the left; scale bar p 0.25 mm. D, Seed coat surface, antiraphal face; scale bar p 100 mm. E–H, Ammannia. E, Ammannia coccinea
Rottb., wetted seed with evaginated epidermal cell trichomes, antiraphal face; scale bar p 100 mm. F, Ammannia coccinea, raphal face,
aerenchymatous float tissue intact; scale bar p100 mm. G, Ammannia erecta (Guill. & Perr.) S.A.Graham & Gandhi, raphal face, float tissue
collapsed; scale bar p 100 mm. H, Ammannia coccinea, one fully and one partially evaginated tuberculate epidermal cell trichome; scale bar p
25 mm. (A from Rios 212; B–D from Breedlove 38133, MICH; E, F, H from Graham 700, MICH; G from Binuyo 172.)

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY 209

Fig. 5 A–C, Capuronia benoistii. A, Antiraphal face; scale bar p 0.5 mm. B, Raphal face; scale bar p 0.5 mm. C, Seed coat surface,
antiraphal face; scale bar p 100 mm. D–G, Crenea. D, Crenea maritima Aubl., oblique view, raphal face to the left; scale bar p 0.5 mm. E,
Crenea patentinervis (Koehne) Standley, antiraphal view with evaginated epidermal cell trichomes; scale bar p 0.5 mm. F, Crenea maritima,
seed coat surface at boundary between raphal (top) and antiraphal (bottom) faces; scale bar p 100 mm. G, Crenea patentinervis, evaginated
epidermal cell trichomes, antiraphal face; scale bar p 50 mm. (A–C from Chauvet 261; D, F from de la Cruz 3301; E, G from Gentry & Juncosa
3854.)

subglobose, apex depressed, (2) 3–5 locular; placenta nonstip-
itate, subglobose, lobes protruding into locules; septa broad,
narrowed slightly distally, ceasing near apex of ovary, placenta
appearing free-basal at maturity; ovules ∼30⫹.
Fruits. Berrylike, 3.5–6.5 mm # 4–6 mm, globose, green
to brown; pericarp apparently thick initially, at full maturity
the inner cell layers and septa forming abundant mucilage into
the locules, on drying the mucilage shrinking closely around
the seeds to a hardened, shiny covering; fruits completely en-
closed in brittle floral tube, indehiscent.
Seeds. Typically 5–8 per berry, 2.9–3.2 mm # 2.3–2.7
mm, golden brown, attached to apical rounded surface of pla-
centa, embedded in pulp; seed shape strongly bilateral, raphal
and antiraphal faces flattened, raphe conspicuous on seed sur-
face; seed outline obovate to cordiform, distinctly narrowed
basally with elongate micropyle, apex obtuse to retuse; seed

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210 INTERNATIONAL JOURNAL OF PLANT SCIENCES
surface smooth, raphal face reticulate toward apex and along
margin, cells polygonal, raphal area tending to scalariform,
cells larger, rectangular, elongate, antiraphal face reticulate to-
ward apex, scalariform toward base in central area, cells po-
lygonal to rectangular and elongate, anticlinal walls of both
surfaces not raised, periclinal walls of antiraphal surface finely
striated.
Epidermal cell internal trichomes. Absent.
Crenea Aubl.
Two species; subshrubs to small trees, flowers actinomor-
phic, hermaphroditic; coastal northern South America, Ec-
uador to Bahia, Brazil; tidal flats, mangroves. Figure 5D–5G.
Ovary. Superior, sessile, globose, 4 (5) locular; placenta
sessile to briefly stipitate, stipe less than 0.2 mm, lobes pro-
truding into locules, ovules developing in tight clusters around
central meristematic points on the lobes, a slender sterile distal
axile placental extension sometimes persisting from ovary
apex; septa ceasing at or just above level of placenta, placenta
free-basal at maturity; ovules 100⫹.
Fruits. Capsules, 5–7 mm # 5–7 mm, globose, red; wall
moderately thick; fruits exceeding floral tube at maturity, ir-
regularly splitting.
Seeds. Approximately 500 per capsule, densely packed,
overlapping, 5.0 mm # 0.3–0.5 mm, fragile, bicolored, raphal
face white or clear, antiraphal face red-brown; seed shape
slightly bilateral, fusiform or narrowly obovoid, raphal face
slightly convex, with aerenchymatous float tissue, flattened to
concave after collapse of float tissue, raphe inconspicuous on
seed surface, antiraphal face convex; seed outline narrowly
oblong, falcate, or clavate, one or two tailed, apex sometimes
slightly enlarged or cucullate, rolled toward raphal face; seed
surface smooth, cells of float tissue thin walled, reticulate, ra-
phal face reticulate, cells polygonal, antiraphal face reticulate
to scalariform, cells rectangular, short, in regular rows toward
apex, more irregular toward base and sides, anticlinal walls
straight, slightly raised, periclinal walls smooth, flat.
Epidermal cell internal trichomes. Present on antiraphal
surface, straight, smooth, without tubercles or ribs, ∼90–95
mm.
Cuphea P.Browne
Approximately 240–250 species, annual and perennial herbs
and shrubs, flowers zygomorphic, bilateral, hermaphroditic;
tropics, subtropics and temperate regions of the Americas; di-
verse wet to dry habitats. Figures 1D, 6, 7, 18A, 18B.
Ovary. Superior, sessile, ovoid to oblong, subtended ex-
ternally by a unilateral basally attached nectariferous organ;
ovary unequally 2 locular, dorsal locule larger, fertile, ventral
locule smaller, fertile or sterile; placenta broad based, sessile
or very briefly stipitate, elongate, narrowly ovoid to oblong,
lobes protruding into the locules; septa vestigial, reduced to
two thin septal threads attaching placenta to ovary wall near
apex of ovary, placenta an elongate axis at maturity; ovules
2–100⫹.
Fruits. Capsules, size varying with species, ∼2/3–3/4 the
length of floral tube, ovoid-oblong, green turning golden; wall
thin, smooth, translucent; fruits included in floral tube at ma-
turity, dehiscence by a single longitudinal slit along dorsal side
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of capsule and a complimentary longitudinal slit in dorsal side
of floral tube, placenta with immature green seeds becoming
erect and emergent through the paired openings, septal threads
ultimately broken on erection of the placenta.
Seeds. Numbering (2) 3–100⫹, typically less than 20 per
capsule, 0.5–3.3 mm # 0.3–3.0 mm, golden brown, brown,
spotted, or purple-black, alternating and overlapping in two
to approximately seven irregular rows per locule; seed shape
slightly to strongly bilateral, subglobose, obovoid, lenticular,
or flattened; seed outline suborbicular, orbicular, cordiform,
obtriangular, obovate, or oblong, apex rounded, truncate, ca-
runculate, or slightly to deeply retuse, raphe conspicuous on
raphal face for length of seed in some species, inconspicuous
on seed surface in others, chalazal area small to prominently
enlarged on raphal face, on both raphal and antiraphal faces,
or not enlarged; seed margin thick and rounded to distinctly
marginate, margin narrowly extended, thinned, encircling,
thus “winged”; seed surface smooth, aveolate, scalariform,
finely to coarsely reticulate, tuberculate (Cuphea circaeoides
Smith) or coarsely wrinkled (Cuphea racemosa [L.f.] Spreng.),
anticlinal walls angular, hexagonal, or linear-rectangular,
raised or not (collinate in young Cuphea micropetala Kunth),
periclinal walls flat or depressed, smooth or striate.
Epidermal cell internal trichomes. Present, helical,
smooth, up to 2 mm long on seeds 3 mm long.
Decodon J.F.Gemel.
One species, Decodon verticillatus (L.) Elliott; perennial
clonal herbs, flowers actinomorphic, tristylic; eastern North
America; lake margins, other wet places. Figures 8A, 8B, 18A.
Ovary. Superior, stipitate with stipe obscured by nectaries
at base, ovary broadly conical to subglobose, 3 (4) locular;
placenta broad based, nonstipitate, elongate, sturdy, with ex-
panded rounded lobes at midlevel in the ovary protruding
slightly into locules; septa sturdy, extending from base to apex
of ovary; placenta an elongate axis at maturity; ovules ∼30⫹.
Fruits. Capsules, 4.5–8.0 mm # 4.5–8.0 mm, globose,
dark brown; wall moderately thick, somewhat indurate; fruits
included to scarcely exceeding floral tube at maturity, dehis-
cence loculicidal.
Seeds. Twenty to 30 per capsule, 1.0–1.9 mm long # 1.2–
2.0 mm wide # 1.0–1.8 mm thick, brown to golden brown,
sometimes with antiraphal germination valve darker, seeds
covering placental lobes, oriented in sunburst arrays; seed
shape obpyramidal, 3–6 faceted, faces sometimes indistinct,
shape often distorted, mostly with three sharp corners and two
rounded corners, raphal face flattened or 2 faceted, raphe cen-
trally positioned, inconspicuous on seed surface, antiraphal
face often 3 or 4 faceted, slightly to strongly convex, area of
germination valve ∼0.8–1.3 mm # 0.6–1.3 mm, cells of valve
raised when fresh and collapsed and darker when desiccated,
occupying ∼2/3 of face, seed outline broadly to narrowly ob-
triangular, apex rounded, two rounded margins of seed slightly
extended as spongy “incipient wings,” wings not apparent ex-
ternally, visible in anatomical transverse section; seed surface
smooth, cell pattern frequently obscure, irregularly reticulate,
cells polygonal, cell pattern of germination valve reticulate to
scalariform, cells polygonal to square or short-rectangular, an-
ticlinal walls not raised, periclinal walls flat.
 Fig. 6 Cuphea. A, B, Cuphea carthagenensis (Jacq.) Macbr. A. Raphal face; scale bar p 1 mm. B, Antiraphal face; scale bar p 1 mm. C,
Cuphea arenarioides A.St.-Hil., raphal face; scale bar p 0.2 mm. D, E, Cuphea micrantha Kunth. D, Antiraphal face; scale bar p 1 mm. E,
Raphal face; scale bar p 1 mm. F, Cuphea urbaniana Koehne, raphal face; scale bar p 0.5 mm. G, H, Cuphea racemosa (L.f.) Spreng. G,
Antiraphal face; scale bar p 0.2 mm. H, Raphal face; scale bar p 0.2 mm. I, Cuphea melvilla Lindl., antiraphal face; scale bar p 1 mm. (A,
B from Graham 442, MICH; C from Reidel., NY; D, E from Zanoni 41271; F from Krummrow 2115; G, H from Boom 760; I from Cavalcanti
et al. 423.)

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 Fig. 7 Cuphea. A–C, E, Cuphea mimuloides Cham. & Schltdl. A, Antiraphal face; scale bar p 0.2 mm. B, Raphal face; scale bar p 0.2
mm. C, Seed coat surface, antiraphal face; note round base of inverted trichome visible on cell walls; scale bar p 50 mm. E, Antiraphal face
with some evaginated trichomes; scale bar p 0.2 mm. D, Cuphea racemosa, seed coat surface, antiraphal face; scale bar p 50 mm. F, G, Cuphea
viscosa Rose. F, Antiraphal face; scale bar p 1 mm. G, Raphal face; scale bar p 1 mm. H, I, Cuphea palustris Koehne. H, Antiraphal face;
scale bar p 1 mm. I, Raphal face; scale bar p 1 mm. J, Cuphea rusbyi Lourt., raphal face; scale bar p 1 mm. (A–C, E from Stevens 19512;
D from Zanoni 41271; F, G from Graham 713; H, I from Graham 804; J from Rusby 117a, US.)

212

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 Fig. 8 A, B, Decodon verticillatus. A, Raphal face, two facets shown; scale bar p 0.5 mm. B, Antiraphal face, arrow at area of germination
valve; scale bar p 0.5 mm. C–G, Didiplis diandra (DC) A.W. Wood. C, Seed coat surface, antiraphal face; scale bar p 50 mm. D, Antiraphal
face; scale bar p 0.2 mm. E, Raphal face, primarily to the left; scale bar p 0.2 mm. F, Germinating seed, radicle emergent at raphal/antiraphal
face boundary; scale bar p 0.2 mm. G, Wetted seed with evaginated straight epidermal cell trichomes, antiraphal face; scale bar p 0.2 mm.
H–J, Diplusodon. H, Diplusodon glaucescens DC., raphal face, prominent wing present; scale bar p 1 mm. I, Diplusodon uninervius Koehne,
raphal face, narrow wing present; scale bar p 1 mm. J, Diplusodon glaucescens, seed coat surface, raphal face, wing to the left, seed body to
the right; scale bar p 0.2 mm. (A from Graham 917; B from Graham 578; C, D, G from Webb s.n.; E from Kellogg s.n.; F, Wooten s.n.; H,
J from Graham 874; I from Cavalcanti et al. 403, CEN, photo with permission of T. Cavalcanti.)

213

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214 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Epidermal cell internal trichomes. Absent.
Didiplis J.F.Gmel.
One species, Didiplis diandra A.W.Wood; aquatic or am-
phibious herbs, flowers actinomorphic, hermaphroditic; en-
demic to eastern United States; ponds, slow-moving streams,
and moist margins, shallow ephemeral waters. Figures 2E, 8C–
8G.
Ovary. Superior, sessile, globose, 2 locular; placenta non-
stipitate, globose, placental lobes protruding into locules; septa
thin, ceasing shortly above the placenta, placenta free-basal at
maturity; ovules 25 to ∼50.
Fruits. Capsules, 1.0–1.7 mm # 1.3–2.2 mm, subglobose,
light brown; wall smooth, thin, translucent; fruits exceeding
floral tube at maturity, irregularly splitting.
Seeds. Twenty-five to 45 per capsule, 0.6–0.9 mm # 0.2–
0.3 mm, brown, totally covering placenta; seed shape bilateral,
narrowly elongate obovoid, raphal face flattened to slightly
convex, antiraphal face strongly convex, raphe inconspicuous
on seed surface; seed outline narrowly obovate, apex rounded
to slightly cucullate; seed surface smooth, raphal face irregu-
larly, coarsely reticulate, cells rectangular, crumpled, antira-
phal face irregularly reticulate, cells rectangular, smaller than
on raphal face, anticlinal walls sinuous, raised, periclinal walls
smooth, flat.
Epidermal cell internal trichomes. Present, straight,
smooth, tapering to apex, 60–80 mm.
Diplusodon Pohl
Eighty-six species (Cavalcanti 2011), perennial shrubs to
small trees, flowers actinomorphic, hermaphroditic; endemic
to Brazil and neighboring areas of northern Bolivia; open cer-
rados, campos, and gallery forests. Figure 8H–8J.
Ovary. Superior, sessile, subglobose, apex depressed, 2
locular, appearing unilocular; placenta broad based, nonstip-
itate, flattened apically, bipartite, protruding into each locule;
septa lunate, thickened at the ovary wall, thinning from ovary
wall toward the placenta, narrowing and ultimately ceasing
above the placenta, placenta free-basal at maturity; ovules 6–
84 per capsule, average 25.
Fruits. Capsules, 3–11 mm # 2–9 mm, oblong to globose,
red-brown, walls thick, indurated; fruits included or exceeding
floral tube at maturity, septicidally dehiscent.
Seeds. Four to 72 per capsule, average 17, 1.0–4.0
mm # 1.8–3.5 mm, light brown, covering apical surface of
the bilobed placenta; seed shape bilateral, lenticular to dis-
coidal, faces slightly convex or flattened, encircled by narrow
wing; wing 0.1–0.7 mm wide, flattened, thinner than central
seed body, raphe conspicuous on seed surface; seed outline
obovate, orbicular, or elliptical; seed surfaces smooth, raphal
face reticulate on seed body, cells of both faces polygonal or
obscure, cells of wing aveolate.
Epidermal cell internal trichomes. Present, straight, ∼800–
1200 mm (T. Cavalcanti, personal communication).
Duabanga Buch.-Ham.
Two or three species; tall trees, flowers actinomorphic, her-
maphroditic; southeastern India, southern China, Malayasia,
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Java, Borneo, Philippine Islands, Celebes, Moluccas, and New
Guinea; subtropical submontane humid forests, swamps, river
margins. Figure 9A, 9B.
Ovary. Semi-inferior, raised toward a superior position
by late growth of upper portions of the ovary, subglobose, 4–
6 (–9) locular; placenta nonstipitate, subglobose, hollow, pla-
cental lobes extensively protruding into the locules; septa thin,
ceasing at ovary apex; placenta axile at lower levels, parietal
at upper levels, ultimately appearing free-basal; ovules 100⫹.
Fruits. Capsules, 1–3.7 cm # 0.8–3.5 cm, subtended by
horizontally spreading sepals, fully exposed at maturity,
broadly globose-ovoid, apex slightly flattened; wall thick,
green, shining, smooth, turning brown, becoming indurated,
woodlike; fruits loculicidally dehiscent.
Seeds. More than 100 per capsule, 4–5.5 mm # 0.5 mm,
light to medium brown, fragile, densely covering rounded en-
larged placental lobes; seed shape slightly bilateral to fusiform
or spindle shaped, 2 tailed, tails 2–2.5 mm long, cells of the
tails large, thin walled, aerenchymatous, central seed body 1
mm long, micropyle elongate, raphe inconspicuous on seed
surface; seed outline narrowly elongate, seed surface scalari-
form, cells short- to long-rectangular, anticlinal walls highly
sinuous, periclinal walls flat, finely pebbled.
Epidermal cell internal trichomes. Absent.
Galpinia N.E.Br.
One species, Galpinia transvaalica N.E.Br.; small trees, flow-
ers actinomorphic, hermaphroditic; southeastern Africa;
woods, thickets, savanna. Figures 9C, 9D, 18A.
Ovary. Superior, stipitate with stipe obscured by nectaries,
ovary subglobose, 2 locular; placenta nonstipitate or stipe less
than 0.5 mm, subglobose, placental lobes protruding into the
locules; septa tapered distally, ceasing at ovary apex; placenta
free-basal at maturity; ovules ∼25–30.
Fruits. Capsules, 3.0–4.5 mm # 2.5–5.0 mm, globose,
pale; wall thin, smooth, translucent; fruits exceeding floral tube
at maturity, irregularly splitting.
Seeds. Three to 10 per capsule, 3.0–4.5 mm # 2.5–4.0
mm, brown, shining, covering surface of the placenta; seed
shape bilateral, lenticular or discoidal, slightly concave-con-
vex, encircled by a broad thin wing 0.3–1.0 mm wide, wing
interrupted at the micropyle, raphe conspicuous on seed sur-
face; seed outline orbicular, seed body orbicular to narrowly
obovoid, ∼1.5–2 mm # 1–1.5 mm; seed surface smooth, ra-
phal and antiraphal surfaces of seed body irregularly reticulate,
cell shapes obscure or polygonal, short-rectangular bordering
the raphe, anticlinal walls slightly raised, cells of wing collinate
or polygonal.
Epidermal cell internal trichomes. Absent.
Ginoria Jacq. (including Haitia Urb., Graham 2010)
Thirteen species; shrubs or trees, flowers actinomorphic, her-
maphroditic; Mexico, Cuba, Hispaniola, Puerto Rico, and Vir-
gin Islands; dry coastal thickets, submontane forests, stream
margins, frequently on serpentine and calcium-rich soils. Fig-
ures 9E–9G, 10A–10D.
Ovary. Superior, sessile, globose to depressed terminally,
2–6 sulcate, 2–6 locular; placenta nonstipitate, ovoid to glo-
bose, outer face of each lobe somewhat flattened, only slightly
 Fig. 9 A, B, Duabanga grandiflora Walp. A, Antiraphal face, central seed body flanked by elongated fragile tails; scale bar p 1 mm. B,
Seed body, anticlinal cell walls highly sinuous; scale bar p 0.2 mm. C, D, Galpinia transvaalica. C, Antiraphal face, convex central seed body
surrounded by wing, interrupted at micropyle/hilum area; scale bar p 1 mm. D, Raphal face, raphe central in concavity; scale bar p 1 mm.
E–G, Ginoria. E, F, Ginoria nudiflora (Hemsl.) Koehne. E, Seed coat surface, antiraphal face; scale bar p 100 mm. F, Seed, raphal face; scale
bar p 0.5 mm. G, Ginoria americana Jacq., raphal face; scale bar p 0.25 mm. (A, B from Maxwell 94–515, CMU; C from Balsinhas 3263,
PRE; D from Pretoria Bot. Institute 238; E, F from Gutiérrez B. 3608, XAL; G from Britton 6742, US.)

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216 INTERNATIONAL JOURNAL OF PLANT SCIENCES

Fig. 10 A, Ginoria callosa O.C.Schmidt, raphal face; scale bar p 0.25 mm. B, Ginoria glabra Griseb., antiraphal face; scale bar p 0.25
mm. C, D, Ginoria pulchra Ekman & O.C.Schmidt. C, Wetted seed with evaginated epidermal cell trichomes, antiraphal face; scale bar p 0.25
mm. D, Evaginated straight epidermal cell trichomes; scale bar p 25 mm. E–G, Heimia salicifolia Link. E, Antiraphal face; scale bar p 0.25
mm. F, Raphal face; scale bar p 0.2 mm. G, Evaginated straight epidermal cell trichomes; scale bar p 25 mm. (A from Ekman 4870; B from
Fairchild Garden, cultivated; C, D from Ekman s.n., S; E–G from Graham 141, MICH.)

protruding into locules; septa narrow, thin, ceasing above the
placenta, placenta free-basal at maturity; ovules 100⫹.
Fruits. Capsules, 1–7 mm # 2–9 mm, globose or ellipsoid,
brown or wine-red; wall moderately thick; fruits included to
exceeding floral tube at maturity, septicidally dehiscent, valves
entire or bipartite at apex.
Seeds. More than 50, 0.5–2.5 mm # 0.1–0.6 mm, golden
to light brown, covering surface of placental lobes; seed shapes,
outlines, and surface patterns diverse, seed shape mostly
slightly to strongly bilateral, narrowly obovoid, apex cucullate
or not, convex-concave, raphe not conspicuous on seed sur-
face, in Ginoria americana Jacq. prismatic with three or four

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY
faces, in Ginoria nudiflora (Hemsl.) Koehne and Ginoria rohrii
(Vahl) Koehne narrowly fusiform and fragile; seed outline ob-
ovate, oblong, elliptic, or subrectangular; seed surface smooth,
cells irregularly reticulate, polygonal or cells linearly arranged,
anticlinal walls typically slightly raised, periclinal walls flat,
or slightly depressed, in G. nudiflora anticlinal walls sinuous
on raphal face.
Epidermal cell internal trichomes. Present, straight,
smooth, ∼175 mm.
Heimia Link.
Three species; shrubs, flowers actinomorphic, hermaphro-
ditic; Texas to Argentina; streamsides, lower montane slopes.
Figure 10E–10G.
Ovary. Superior, sessile, turbinate to subglobose, apex
slightly depressed, 4 (5) sulcate, 4 (5) locular; placenta non-
stipitate or with stipe less than 0.5 mm, elongate ovoid to
subglobose, placental lobes strongly protruding into the loc-
ules; septa ceasing near ovary apex; placenta an elongate axis
at maturity; ovules 100⫹.
Fruits. Capsules, 2–5 mm # 2.3–5.5 mm, subglobose,
brownish green or brown; wall moderately thick; fruits in-
cluded in floral tube at maturity, loculicidally dehiscent.
Seeds. More than 50, 0.5–1.0 mm # 0.3–0.8 mm, golden
to light brown, densely covering surfaces of placental lobes;
seed shape bilateral, narrowly obovoid, raphal face 2 faceted,
raised or flattened along the raphe, antiraphal face convex, 1–
3 faceted, seed apex flattened with smooth corners or cucullate,
raphe inconspicuous on seed surface; seed outline narrowly
obtriangular or oblong, apex polygonal; seed surface smooth,
cells on both faces reticulate, irregularly rectangular, elongate,
anticlinal walls straight or sinuous, not or slightly raised, per-
iclinal wall smooth, flat.
Epidermal cell internal trichomes. Present, straight,
smooth or with raised striae basally, 70–80 mm.
Koehneria S.A.Graham, H.Tobe & P.Baas.
One species, Koehneria madagascariensis (Baker)
S.A.Graham et al.; shrubs, flowers actinomorphic, hermaph-
roditic; endemic to Madagascar; semiarid savannas. Figure
11A–11C.
Ovary. Superior, stipitate with stipe 1 mm, ovary globose,
2–4 locular; placenta nonstipitate, ovoid to globose, placental
lobes strongly protruding into the locules; septa ceasing at
ovary apex; placenta an elongate axis at maturity; ovules
100⫹.
Fruits. Capsules, 4.5–5.0 mm # 4.3–6.0 mm, oblong to
globose, brown; wall thin, smooth, bearing sessile, globose,
black resin-filled trichomes; fruits included to slightly exceed-
ing floral tube at maturity, subtended by reflexed sepals, sep-
ticidally dehiscent.
Seeds. Approximately 10–30, 1.3–1.7 mm # 1–1.2 mm,
golden to deep brown, covering surface of placenta distal to
sterile stipe; seed shape slightly bilateral, approximately cu-
neiform, 3–5 faceted, raphal face flattened or 2 faceted, raphe
raised on seed surface, antiraphal face 2 or 3 faceted or margins
rounded and face convex; seed outline narrowly to broadly
cuneate, apex flat, polygonal in outline; seed surface smooth,
cells of raphal face irregularly linearly arranged, short-rect-
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217
angular to polygonal, more elongate over the raphe, cells of
antiraphal surface more linearly arranged, short-rectangular,
anticlinal walls slightly raised, periclinal walls flat to depressed,
finely striated.
Epidermal cell internal trichomes. Present, helical, 300
mm.
Lafoensia Vand.
Five or six species; shrubs or trees, flowers actinomorphic,
hermaphroditic; southern Mexico to southern Brazil; cerrados
and low montane forests. Figures 1A, 11D–11F, 18A, 18C.
Ovary. Superior, stipitate with stipe half the length of
ovary to longer than ovary, ovary turbinate or globose, wall
smooth or encircled at the equator by short vertical depres-
sions, 2 or 4 locular, appearing unilocular; placenta nonstip-
itate, initially globose or subglobose, ultimately apically flat-
tened, disciform; septa vestigial, almost absent, placenta
free-basal at maturity; ovules 100⫹.
Fruits. Capsules, 1.0–6.5 cm # 2.5–8.0 cm, subglobose
to obovoid or turbinate, brown to reddish; wall thick, smooth
or with depressions like the ovary surface, initially leathery,
ultimately dry, indurate, woodlike; fruits included in floral
tube, floral tube falling away with age, fruit loculicidally de-
hiscent, 2 or 4 valved.
Seeds. More than 100, 6.5–35 mm # 8–15 mm, reddish,
yellow, or brown, covering flattened apical surface and sides
of the placenta; seed shape bilateral, discoidal, encircled by a
thin glassy wing 1–5 mm wide, raphe scarcely raised on seed
surface; seed outline subcircular to oblong or rectangular, seed
body oblong to obovoid, micropylar extension at base of seed
body extending to base of wing; seed surface smooth, reticulate
to scalariform or obscure, cells of both faces variably polyg-
onal to rectangular-collinate, cells of wing highly
mucilaginous.
Epidermal cell internal trichomes. Present, helical, 600
mm.
Lagerstroemia L.
Approximately 56 species; shrubs and trees, flowers acti-
nomorphic, hermaphroditic; India, China, Southeast Asia, Ma-
laysia, New Guinea, northern Australia, Philippines north to
Japan; riverine forests, montane and secondary forests. Figures
1E, 12D, 12G, 18A.
Ovary. Superior, sessile, slightly imbedded in the recep-
tacle, broadly ovoid, (3–) 6 locular; placenta nonstipitate, nar-
rowly ovoid, glandular, axis slender, placental lobes scarcely
intruding into locules; septa sturdy, remaining broad above the
placenta, ceasing near apex of ovary, secondary septal protru-
sions extending from midpoint between the major septa into
locules less than one-fifth distance to central axis; placenta an
elongate axis at maturity; ovules ∼8–100.
Fruits. Capsules, typically ∼1.5–2.0 cm # 1.0–1.7 cm,
ranging from ∼1–2.5 cm # 1–1.5 cm (Lagerstroemia parvi-
flora Roxb.) to 3.5 cm # 2.5 cm (Lagerstroemia costa-dra-
conis Furtado & Srisuko), oblong, elliptic, ovoid, or globose,
light to deep brown, rarely black; wall thick, leathery, becom-
ing indurated, woodlike; fruit exceeding floral tube at maturity,
loculicidally dehiscent.
Seeds. Eight to ∼72⫹ per capsule, seed including unilateral
218 INTERNATIONAL JOURNAL OF PLANT SCIENCES

Fig. 11 A–C, Koehneria madagascariensis. A, Antiraphal face, two faceted, base torn; scale bar p 0.2 mm. B, Raphal face; scale bar p
0.2 mm. C, Seed coat surface, antiraphal face; scale bar p 50 mm. D–F, Lafoensia nummularifolia A. St.-Hil. D, Seed coat surface, seed body,
raphal face; scale bar p 0.5 mm. E, Raphal face, encircling wing, central seed body, elongate micropyle; scale bar p 2 mm. F, Wetted seed,
evaginated helical trichomes on surface of the wing; scale bar p 1 mm. (A from Lowry et al. 5922; B from van der Werff & MacPherson
13595; D–F from Graham 912.)

flattened wing, ∼3.0–6.0 mm # 4.0–16.0 mm, seed body only Lawsonia L.

∼3.0–6.0 mm # 1.5–2.5 mm, seeds arranged in two longi-
tudinal rows in each locule on either side of the secondary
septa, attached to placenta at the pointed “base,” wings ex-
tending distally toward ovary apex, flattened against the septa;
seed shape of seed body obpyramidal or nearly so; outline of
wing subrectangular, oblong, or slightly falcate, wing extend-
ing from raphal face, thickest on the top long margin, thinning
basally, raphe not conspicuous on seed surface; seed surface
smooth on seed body and wing, cells mostly irregularly ar-
ranged across the width of seed body, polygonal or collinate,
cells of wing oriented with wing length, polygonal to mostly
short-rectangular, anticlinal walls not raised, periclinal walls
flat or slightly raised.
Epidermal cell internal trichomes. Absent.
One species, Lawsonia inermis L.; shrubs, flowers actino-
morphic, hermaphroditic; Eurasian in origin, widely cultivated
worldwide in subtropical and tropical climates; savannas,
semiarid zones. Figures 12A–12C, 18D.
Ovary. Superior, sessile, subglobose to globose, (2–) 4
locular; placenta stipitate with stipe ∼1/4 to 1/3 length of
ovary, placenta sturdy at base, slender at midovary height with
(2–) 4 semiflattened branched lobes protruding into locules,
fragile placental extension often present above the lobes; septa
thin, narrowed toward apex of capsule, ceasing at ovary apex;
placenta an elongate axis at maturity (also free basal?); ovules
∼70–80, in clusters on branches of the placental lobes.
Fruits. Capsules, 3.5–7 mm # 4.5–8 mm, subglobose,

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY 219

Fig. 12 A–C, Lawsonia inermis. A, Antiraphal face, arrow indicating area of distinctive cellular arrangement suggestive of a germination
valve; scale bar p 1 mm. B, Raphal face, seed slightly immature; scale bar p 1 mm. C, Seed coat surface, seed apex; scale bar p 0.2 mm. D,
G, Lagerstroemia. D, Lagerstroemia speciosa (L.) Pers., seed body to the left, wing extending from raphal face to the right; scale bar p 2 mm.
G, Lagerstroemia indica L., seed coat surface of seed body adjacent to wing; scale bar p 0.2 mm. E, F, H, Lourtella resinosa. E, Antiraphal
face, trifaceted; scale bar p 0.2 mm. F, Raphal face; scale bar p 0.2 mm. H, Seed coat surface, antiraphal face; scale bar p 50 mm. (A–C from
Correll 45915, TEX; D from Fairchild Gardens, cultivated; G from cultivated plant, no voucher; E, F, H from Lopez & Sagastegui 8229.)

mostly wider than long, brown; wall moderately thick, be-
coming dry, brittle, enveloped at base by sepals, exceeding the
floral tube at maturity, irregularly splitting.
Seeds. Approximately 30–50 per capsule, 1.2–2.2 mm #
1.3–2.0 mm, red-gold brown, surface shining; distal one-third
to one-half composed internally of small-celled, dense, spongy
float tissue; seed shape short-obpyramidal, corners rounded,
apex flat to shallowly rounded, broad, 3–6 faceted, base slen-
der, pointed, raphe visible on lower half of raphal surface; seed
outline obtriangular or slightly flattened; seed surface smooth,
pattern irregular, cells of both faces on upper one-third to one-
half of seed large, thin walled, foveolate, cells below smaller,
more irregular in size and arrangement, walls somewhat
obscure.
Epidermal cell internal trichomes. Absent.
Lourtella S.A.Graham, P.Baas, & H.Tobe.
One species, Lourtella resinosa S.A.Graham et al.; shrubs,
flowers actinomorphic, hermaphroditic; northern Peru to
southern Bolivia; dry open woodlands. Figure 12E, 12F, 12H.
Ovary. Superior, sessile, globose, 2 locular; placenta with

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220 INTERNATIONAL JOURNAL OF PLANT SCIENCES
a 0.2-mm sterile stipe, slender ovoid to subglobose, protruding
massively into the locules; septa vestigial, narrow, reduced at
apex of placenta to two threadlike remnants joining placenta
to ovary wall near ovary apex; placenta an elongate axis at
maturity (study material limited, possibly free-basal at matur-
ity); ovules ∼50–60.
Fruits. Capsules, 3–4 mm # 3–4 mm, globose, pale
brown, flushed red with age, thin walled, smooth, papery,
translucent, included in floral tube at maturity, irregularly
splitting.
Seeds. Approximately 30–50 per capsule, 0.8–1 mm #
0.5–0.7 mm, brown, attached over entire placental surface;
seed shape short-obpyramidal, 3–5 faceted, facets shallow,
margins rounded, raphal face slightly convex, raphe conspic-
uous, defining margin of two adjoining facets, antiraphal sur-
face shallowly 1–3 faceted or flattened; seed outline obtrian-
gular, apex flat, base rounded to truncate; seed surface smooth,
cells polygonal to short-rectangular, more or less linearly ar-
ranged, especially on antiraphal face, anticlinal walls straight,
periclinal walls flat.
Epidermal cell internal trichomes. Absent.
Lythrum L. (including Peplis L., Webb 1967)
Approximately 35 species; annual and perennial herbs, flow-
ers actinomorphic, hermaphroditic, some di- or tristylous; tem-
perate Europe, Asia, Americas; wet places, swamps, lake mar-
gins, ditches. Figure 13.
Ovary. Sessile to short-stipitate with stipe partially or fully
surrounded by circular nectary, ovary ovoid to elongate; 2
locular; placenta nonstipitate or stipitate with stipe ∼0.2 mm
(in Lythrum portula [L.] D.A.Webb), placenta mostly an elon-
gate axis, slender ovoid and lightly compressed (L. salicaria,
L. alatum), placental lobes slightly protruding into the locules,
septa thin, typically ceasing near ovary apex (infrequently, in
species formerly in Peplis, the placenta globose and appearing
free-basal at maturity); ovules typically 30–100⫹.
Fruits. Capsules, ∼2.0–5.0 mm # 1.5–2.0 mm, oblong
or globose, deep brown or reddish-brown; wall thin; fruits
included or exceeding the floral tube, septicidally dehiscent,
valves bipartite at apex in some species.
Seeds. Approximately 30–120 per capsule, 0.4–0.8
mm # 0.3–0.7 mm, golden brown, partly overlapping in two
rows per locule on elongate placentas, crowded and covering
surface of globose placentas; seed shape bilateral, obovoid to
subglobose, narrowly obovoid, or ellipsoid, raphal face flat-
tened or slightly concave, raphe inconspicuous, antiraphal face
convex; seed outline suborbicular, obovate, oblong to elliptic;
seed surface smooth, irregularly reticulate, cells rectangular,
cells of raphal face larger and more elongated than on anti-
raphal face, anticlinal walls straight, raised, periclinal walls
flat or slightly collapsed, in some species finely striated width-
wise.
Epidermal cell internal trichomes. Present, emerging at
one end of the periclinal wall of the rectangular epidermal
cells, straight, smooth (L. alatum Pursh, L. californicum Torr.
& Gray, L. maritimum Kunth, L. salicaria L.) or shallowly to
deeply ribbed (L. borsysthenicum [Schrank] Litv., L. portula
[L.] D.A. Webb, L. volgense D.A. Webb), 60–80 mm.
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Pehria Sprague
One species, Pehria compacta (Rusby) Sprague; shrubs or
small trees, flowers actinomoprhic, hermaphroditic; Hondu-
ras, Nicaragua, Venezuela, Colombia, in secondary vegetation,
roadsides, pastures, margins of low- to midelevation forests.
Figure 14A–14C, 14F.
Ovary. Superior, stipitate with stipe 1–2 mm, oblong to
subglobose, 2 locular; placenta nonstipitate or with stipe to
0.2 mm, oblong to globose, placental lobes protruding exten-
sively into locules; septa very narrow (∼0.4 mm wide at base),
tapering distally, ceasing at apex of placenta, placenta free-
basal at maturity; ovules 100⫹.
Fruits. Capsules, ∼6.0 mm # 5.0 mm, subglobose, red-
dish-brown; walls thick, indurate, fruits included in floral tube
at maturity, basically loculicidal, sometimes initially circum-
scissile around base of sturdy style and then loculicidally de-
hiscent, valves bipartite at apex.
Seeds. More than 50 per capsule, 0.7–0.9 mm # 0.3–0.5
mm, brown, covering surface of placental lobes; seed shape
bilateral, narrowly obovoid, raphal face slightly 2 faceted or
flattened, raphe inconspicuous on seed surface or defining mar-
gin of adjacent facets, antiraphal face convex; seed outline
narrowly obovate, base pointed to obtuse, apex rounded; seed
surface smooth, scalariform to reticulate on both surfaces, cells
rectangular, larger on raphal surface, anticlinal walls straight,
slightly raised, periclinal walls smooth to slightly depressed.
Epidermal cell internal trichomes. Present, straight, com-
pressed, almost bifacial, finely striated by minute elongated
tubercles, 50–100 mm.
Pemphis J.R.Forst. & G.Forst.
One species, Pemphis acidula J.R.Forst. & G.Forst.; shrubs,
flowers actinomorphic, hermaphroditic; Indian and Pacific
Oceans from East Africa to Marshall Islands and north to
Ryukyu Islands; sandy marine beaches, thickets along coastal
inlets. Figure 14D, 14E.
Ovary. Superior, sessile, globose, apex slightly depressed,
3 or 4 locular; placenta nonstipitate or with stipe less than 0.2
mm, subglobose, lobes slightly protruding into locules, apex
slightly flattened; septa ceasing abruptly above placenta at mid-
level of ovary, placenta free-basal at maturity; ovules ∼20⫹.
Fruits. Capsules, 6.0–6.5 mm # 5–6 mm, oblong-ellip-
tical, reddish brown to brown; wall thick, indurate; fruits
slightly exceeding floral tube at maturity, dehiscence
circumscissile.
Seeds. Fifteen to 20 per capsule, 2.5–3.0 mm # 1.7–2.0
mm, golden to reddish brown, shining, crowded in capsule,
attached on flattened placental apex and sides; seed shape
highly irregularly obpyramidal, variously 3–6 faceted, seed
body central, narrowly obovate, ∼1.7 mm # 1.2 mm, nar-
rowly tapering to base, apex flattened or nearly so, seed in-
ternally surrounded and topped by angular “wings” of densely
packed small-celled spongy tissue, raphe conspicuous on seed
surface; seed outline irregularly obtriangular to oblong; seed
surface smooth, cells of raphal face irregularly reticulate, small
rectangular to polygonal, parallel along the raphe, cells of an-
tiraphal face reticulate, narrower, more elongate-rectangular
than cells of raphal face, or polygonal, or obscure.
Epidermal cell internal trichomes. Absent.
 Fig. 13 Lythrum. A–C, G, Lythrum borysthenicum Rouy & E.G.Camus. A, Antiraphal face; scale bar p 0.2 mm. B, Raphal face; scale
bar p 0.2 mm. C, Evaginated straight epidermal cell trichomes, antiraphal face; scale bar p 20 mm. G, Antiraphal face, periclinal walls striate,
trichomes initiating evagination on the right; scale bar p 20 mm. D, E, Lythrum salicaria L. D, Oblique view, raphal face to the left, antiraphal
face to the right; scale bar p 0.2 mm. E, Seed coat surface, antiraphal face; scale bar p 50 mm. F, H–J, Lythrum californicum Torr. & Gray.
F, Seed coat surface, antiraphal face; scale bar p 50 mm. H, Oblique view, raphal face to the left, antiraphal face to the right; scale bar p 0.2
mm. I, Wetted seed with evaginated straight epidermal cell trichomes, antiraphal face; scale bar p 0.2 mm. J, Evaginated epidermal cell trichomes,
antiraphal face; scale bar p 25 mm. (A–C, G fromTzvelev & Petchenynk s.n.; D, E from Whetstone 6608; F, H–J from Graham 701.)

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 Fig. 14 A–C, F, Pehria compacta. A, Oblique view, antiraphal face to the left, raphal face to the right; scale bar p 0.2 mm. B, Wetted seed
with evaginated straight epidermal cell trichomes, antiraphal face; scale bar p 0.25 mm. C, Evaginated trichome in the center, partially evaginated
trichome in lower right; scale bar p 20 mm. F, Surface of trichome with elongated tubercles; scale bar p 5 mm. D, E, Pemphis acidula. D,
Antiraphal face, arrow indicating distinctive cellular area over the seed body; scale bar p 1 mm. E, Raphal face; scale bar p 1 mm. (A–C, F
from Berry 4023; D from Flynn 6217; E from Vandervelde s.n., no voucher.)

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY
Physocalymma Pohl
One species, Physocalymma scaberrimum Pohl; trees, flow-
ers actinomorphic, hermaphroditic; central South America; dry
forests and savannas. Figure 15A, 15E.
Ovary. Superior, slightly embedded in the receptacle,
subglobose, 4 locular; placenta nonstipitate, subglobose, lobes
protruding into locules; septa narrow, tapered and ceasing
above the placenta, placenta free-basal at maturity; ovules
50⫹.
Fruits. Capsules, 8.0–9.0 mm # 9.0–11.0 mm, subglo-
bose, wall thin, translucent; fruits included in floral tube at
maturity, irregularly splitting.
Seeds. More than 50 per capsule, 3.2–4.2 mm # 2.7–4.0
mm including encircling wing, light brown, partially overlap-
ping on placental surface; seed shape bilateral, discoidal, mar-
gin irregular, seed body 1.5–2.5 mm # 1.5–1.8 mm, slightly
raised above wing surface on antiraphal face, micropyle elon-
gate, 0.5–1 mm, extended through wing, wing thin, 0.8–1 mm
wide, encircling the seed body, raphe visible on seed surface;
seed outline suborbicular, seed body obovate or oblong; seed
surface irregularly reticulate, cells of both faces polygonal to
short-rectangular, vaguely linearly radiating from margin of
seed body across wing surface to wing margin, cells along
length of the raphe more elongate-rectangular, cells of re-
mainder of seed body reticulate, polygonal or obscure.
Epidermal cell internal trichomes. Absent.
Pleurophora D.Don.
Approximately seven species; perennial herbs, flowers zy-
gomorphic, bilateral, hermaphroditic; Brazil, Bolivia, Chile,
Argentina; semiarid open scrub to true desert habitats. Figures
2F, 15B–15D, 15F, 15G.
Ovary. Superior, stipitate with stipe to 5 mm, encircled
by nectary, ovary approximately ovoid, unequally 2 locular,
dorsal locule larger, fertile, ventral locule smaller, fertile or
sterile, dorsal locule slightly to prominently dorsally gibbous;
placenta stipitate with stipe ∼1 mm, placenta elongate, ovoid,
dorsal placental lobe enlarged, nearly filling the locule, ventral
lobe less developed with fewer ovules (sterile in Pleurophora
anomala [St.Hil.] Koehne), scarcely protruding into ventral
locule; septa vestigial, reduced to two short threads at apex
of placenta joining placenta to ovary wall near ovary apex;
placenta an elongate axis at maturity; ovules ∼25–100⫹.
Fruits. Capsules, ∼3.0 mm # 2.0 mm (in Pleurophora
saccocarpa, in species with smaller flowers the capsules pro-
portionally smaller), light brown; wall thin, translucent; fruit
included at maturity, irregularly splitting.
Seeds. Three to ∼100 per capsule, 0.6–0.7 mm # 0.4–
0.5 mm, dark brown-black; seed shape bilateral, obovate to
oblong-ellipsoid, raphal face flattened to slightly convex, raphe
conspicuous on seed surface the length of seed, antiraphal face
convex; seed surface smooth, raphal face reticulate at seed
margin, irregularly reticulate and crumpled at center of the
face, cells rectangular to square, antiraphal face scalariform
to reticulate, cells rectangular, anticlinal walls mostly straight,
slightly raised, periclinal walls smooth, flat to depressed.
Epidermal cell internal trichomes. Present, helical, emerg-
ing at distal end of cells, 120–200 mm.
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223
Punica L.
Two species; shrubs or small trees, flowers actinomorphic,
hermaphroditic; Punica granatum L., pomegranate, probably
native to area of modern Iran, widely cultivated, Punica pro-
topunica Balf.f., endemic to Socotra; semiarid regions. Figure
15H, 15K.
Ovary. Superior to semi-inferior and globose in P. pro-
topunica with carpels in one whorl, 3 locular (Levin 1980) or
5–7 locular (Balfour 1888), semi-inferior to inferior and glo-
bose in P. granatum with two twisted whorls of carpels, an
upper whorl of ∼8 or 9 carpels, superposed on a lower one
of 3 or 4 carpels (but see Sinha and Joshi 1959, p. 43, who
cite up to 3 whorls and up to 15 carpels); septa thick, tough,
extending to apex of ovary; placentation parietal in P. pro-
topunica, ovules covering the bottom and walls of the ovary
(Balfour 1888), placentation in P. granatum axile in the lower
whorl, parietal in upper whorl; ovules 100⫹.
Fruits. Berries, 4.5 cm # 5.0 cm in P. protopunica; ∼5.0–
12.0 cm # 5.0–12.0 cm in P. granatum (cultivated fruits),
subglobose to globose, yellow to red or red-brown, “crown”
of sepals and stamens persistent at the fruit apex; fruit wall
(persistent floral tube plus carpel walls) thick, leathery exter-
nally, spongy internally; indehiscent.
Seeds. More than 100 per berry (in P. granatum more
than 1000 in large cultivated fruits), 9.0–12.0 mm # 7.0–9.5
mm including juice-filled sarcotesta, sarcotesta ruby red, pink,
or yellowish white, ∼2 mm thick, seed body minus sarcotesta
∼4.0–6.5 mm # 3–4 mm in P. granatum, 4.0–4.6 mm # 1.8–
2.0 mm in P. protopunica minus sarcotesta (Miller & Merello
7881 [MO]), seeds attached and embedded in irregularly
twisted, white septal and placental tissue and spongy tissue of
inner ovary wall; seed shape including sarcotesta subobpy-
ramidal 3–6 faceted, apex flattened or slightly convex, base
with circular depression, funicle extending through the sar-
cotesta to inner seed layers (mesotestal layers), micropyle and
hilum obscure, raphe not visible, seed shape minus sarcotesta
ellipsoid, raphal face slightly convex to flattened, antiraphal
face convex, sometimes 2 faceted; seed outline with sarcotesta
oblong to obovoid, minus sarcotesta elliptic to oblong; seed
surface smooth, reticulate, sarcotestal cells hexagonal, anti-
clinal walls not raised, periclinal walls flattened, smooth.
Epidermal cell internal trichomes. Absent.
Rotala L.
Fifty species; aquatic to terrestrial annual herbs, flowers ac-
tinomorphic, hermaphroditic, some distylous; Africa, South-
east Asia, two species in the Americas; ponds, margins of
streams, wet places. Figures 2A–2C, 15I, 15J, 15L.
Ovary. Superior, sessile, globose or oblong, 2–5 locular;
placenta nonstipitate to stipitate with stipe up to one-fourth
length of ovary, placenta globose or oblong, enlarged lobes
slightly protruding into locules; septa thin, narrow, ceasing at
apex of placenta, placenta free-basal at capsule maturity;
ovules 40⫹.
Fruits. Capsules, 0.5–2.5 mm # 0.2–2.0 mm, golden to
light brown or red; walls thin, papery, translucent, finely trans-
versely striated; fruits included to exceeding the floral tube,
septicidally dehiscent.
Seeds. Approximately 30–300 per capsule, 0.3–0.7 mm
 Fig. 15 A, E, Physocalymma scaberrimum. A, Antiraphal face; scale bar p 1 mm. E, Seed coat surface, raphal face; scale bar p 0.2 mm.
B–D, F, G, Pleurophora saccocarpa Koehne. B, Antiraphal face; scale bar p 0.2 mm. C, Raphal face; scale bar p 0.2 mm. D, Wetted seed,
antiraphal face with evaginated helical epidermal cell trichomes; scale bar p 0.2 mm. F, Seed coat surface, antiraphal face; scale bar p 50 mm.
G, Evaginated helical trichomes, antiraphal surface; scale bar p 50 mm. H, K, Punica granatum. H, Raphal face, sarcotesta and outermost
layers of the mesotesta removed; scale bar p 1 mm. K, Sarcotesta surface, raphal face; scale bar p 50 mm. I, J, L, Rotala ramosior (L.) Koehne.
I, Antiraphal face; scale bar p 100 mm. J, Raphal face, aerenchymatous cells collapsed; scale bar p 100 mm. L, Evaginated straight epidermal
cell trichomes, antiraphal face; scale bar p 100 mm. (A, E from Killeen & Perez 6680; B–D, F, G from Krapovickas & Cristobal 12984; H,
K from cultivated fruit; I, L from Graham 1055; J from Graham 438.)

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY 225

Fig. 16 Sonneratia. A, C, Sonneratia caseolaris (L.) Engl. A, Seed; scale bar p 0.5 mm. C, Interior (mesotestal) layers; scale bar p 100
mm. B, D, Sonneratia ovata Backer. B, Seed, most of dark epidermal surface cells fallen away exposing cream-colored interior (mesotesta); scale
bar p 1 mm. D, Epidermal surface, oval to orbicular mucilage cells of the exotesta present; scale bar p 0.2 mm. (A, C from Graham s.n. in
2003; B, D from P. K. Char s.n. in 1990.)

(1.5 in Rotala fimbriata Wight) # 0.2–0.4 mm, brown, golden,
red-brown or red, covering surface of the placenta; seed shape
slightly bilateral, obovoid or ellipsoid to subglobose, raphal
face covered by an arenchymatous float of 2–3 layers of large
thin-walled cells, face slightly concave with collapse of float
tissue, raphe sometimes visible on seed surface, antiraphal face
strongly convex; seed outline suborbicular to obovate, elliptic;
seed surface smooth, raphal face irregularly reticulate, cells
rectangular to square or obscure on the collapsed convex sur-
face, antiraphal face reticulate to scalariform or irregularly
reticulate, cells long-rectangular; anticlinal walls straight,
scarcely or not raised, periclinal walls smooth.
Epidermal cell internal trichomes. Present in most species
(apparently absent in aquatic Rotala mexicana), straight,
smooth, shallowly ribbed or roughly tuberculed, 70–100 mm.
Sonneratia L.f.
Seven species; small to medium-sized trees, flowers actino-
morphic, hermaphroditic; coastal tropical East Africa to Indo-
Malesia, islands of the western Pacific; mangrove swamps,
tidal estuaries, sheltered bays. Figure 16A–16D.
Ovary. Superior to semi-inferior, short stipe obscured by
encircling nectary (in Sonneratia alba Sm.), globose, depressed
terminally, (4–) 15 or 16 (⫺21) locular; placenta subglobose,
thick, fleshy, hollow, apex flattened, enlarged placental lobes
protruding slightly into locules at approximately midlevel of
ovary; septa thin, broad, ceasing above the placenta, placenta
axile at lower levels and parietal at upper levels of ovary,
ultimately appearing free-basal; ovules 100⫹ (to more than
1000).

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226 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Fruits. Berries, 2.5–4.0 cm # 3.0–8.5 cm (excepting Son-
neratia apetala, thin walled, 12–18 mm # 2.3 cm), broadly
adhering to a persistent flattened floral base or sunken into
and partially fused with surrounding floral cup, globose or
depressed globose, sometimes indented around style base,
green, yellow-green, turning brown or grayish; wall thick,
leathery, smooth, shining or dull, nonribbed; fruits well ex-
serted at maturity, indehiscent. (Study material of ovary and
placenta limited.)
Seeds. More than 100 to more than 1000 per fruit, 5–12
mm # 4–10 mm, embedded in acidic pulp, tightly packed over
the surface of the placental lobes in two irregular rows per
locule, corky brown, whitish when epidermis is missing; shape
bilateral, flattened or irregularly angular, raphe extending
along one “arm” of seed, inconspicuous on seed surface; seed
outlines diverse, often U or V shaped; seed surface rough,
thickened, epidermal cells varying in size and shape, drying
red-brown, mucilaginous, when wet, swelling, bulliform, fall-
ing away from inner cell layers of small cubical cells.
Epidermal cell internal trichomes. Absent.
Tetrataxis Hook.f.
One species, Tetrataxis salicifolia (A.Thouars ex Tul.) Baker;
small trees, flowers actinomorphic, hermaphroditic; rare en-
demic of Mauritius; in lower montane wet forest. Figure 17A,
17B.
Ovary. Superior, sessile, broadly ovoid, 4 or 5 locular;
placenta sessile or very short-stipitate, placenta elongate-ovoid,
extending to apex of the ovary, placental lobes enlarged, pro-
truding into the locules; septa thin, narrow, continuing to apex
of ovary; placenta an elongate axis at maturity; ovules 100⫹.
Fruits. Capsules, 6.0–7.0 mm # 7.0–8.0 mm, globose,
brown; wall moderately thick, somewhat indurate; fruit in-
cluded in floral tube at maturity, dehiscence initially Ⳳcircum-
scissile at base of style and then septicidal.
Seeds. More than 50 per capsule, 0.9–1.2 mm # 0.2–0.5
mm, light brown, covering the placental lobes; seed shape bi-
lateral to subcylindric, narrowly obovoid or short-fusiform,
raphal face slightly to strongly concave, raphe inconspicuous
on seed surface, antiraphal face convex, apex frequently cu-
cullate; seed outline oblong, obovate, or elliptic; seed surface
smooth, scalariform or reticulate to irregularly reticulate, cells
of both surfaces rectangular, anticlinal walls straight, slightly
raised, periclinal walls shallowly depressed, smooth.
Epidermal cell internal trichomes. Absent. (Material for
study limited.)
Trapa L.
Three species or more, number of species unsettled; annual
aquatic herbs, flowers actinomorphic, hermaphroditic; Eu-
rope, Africa, Asia, invasive in North America and Australia;
ponds, slow-moving rivers. Figure 17C, 17D.
Ovary. Semi-inferior at anthesis, subsequently becoming
inferior in fruit, approximately obovoid, surrounded by a cor-
onary disc, 2 locular; inner epidermis of ovary wall, placenta,
and funicle glandular, secretory; placenta apical, fertile at
ovary apex, single ovule pendulous from apex in each locule,
one developing, one vestigial; septum complete; ovules 2.
Fruits. Drupe-like, 2–4 horned, fleshy floral cup rhombic,
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turbinate, ∼1.0–3.0 cm long # 1.0–5.0 cm wide (spine tip to
spine tip) # 1.0–3.0 cm thick (Suriyagoda et al. 2006); outer
layers of persistent floral cup evanescent, remaining tissues of
flower and fruit hardened, stony, 2–4 sepals forming conical
to blunt horns, accessory hardened tubercles sometimes pres-
ent on surface; fruit indehiscent.
Seeds. One per fruit, retained in stony floral cup/fruit
structure, seed size ∼1.0–2.0 cm # 1.0–2.0 cm estimated from
fruit dimensions, seed shape approximately obpyramidal or
slightly compressed bilaterally, seed coat cell layers compressed
at embryo maturity, embryo large, starchy. (Living material
not studied.)
Epidermal cell inverted trichomes. Absent.
Woodfordia Salisb.
Two species; shrubs or small trees; flowers incipiently zy-
gomorphic; Africa, Madagascar, Pakistan to China and In-
donesia, semixeric areas, exposed slopes. Figure 17E–17G.
Ovary. Superior, sessile, slightly bilateral, narrowly ovoid-
oblong, 2 locular, dorsal locule slightly larger than the ventral;
placenta sessile to very short-stipitate, elongate, ellipsoid, or
narrowly ovoid, extending to apex of ovary, lobes protruding
extensively into the locules; septa narrow, ceasing at apex of
placenta at the ovary apex; placenta elongate at capsule ma-
turity; ovules 100⫹.
Fruits. Capsules, 7.5 mm # 4.0 mm, narrowly ovoid, thin
walled, papery, translucent; fruits included in persistent, dis-
tended floral tube at maturity, irregularly splitting along one
or more longitudinal lines of the capsule wall and floral tube.
Seeds. More than 50, 0.7–1.0 mm # 0.2–0.3 mm, golden
brown, covering entire elongate placenta except short basal
sterile stipe; seed shape slightly bilateral, narrowly obpyram-
idal or subcylindric, base rounded, obtuse, or truncate, apex
flattened, raphal face flattened to shallowly concave, antira-
phal face convex, apex sometimes slightly cucullate, raphe
raised and conspicuous over length of the seed surface; seed
outline narrowly obovate to oblong; seed surface smooth, sca-
lariform, irregularly reticulate toward the base, cells rectan-
gular, anticlinal walls straight to slightly sinuous, raised, peri-
clinal walls shallowly depressed, longitudinally striate.
Epidermal cell internal trichomes. Present, helical, 70 mm
long.
Diversity of Ovary, Fruit, and Seed Characters
The Ovary
Superior ovaries were long considered a major diagnostic
feature of the Lythraceae. This changed with the inclusion of
the Duabangaceae, Sonneratiaceae, Punicaceae, and Trapaceae
in an expanded Lythraceae (Graham et al. 2005). A superior
ovary still dominates in the Lythraceae sensu lato, but in Dua-
banga the ovary is semi-inferior (Jayaweera 1967; Tobe et al.
1998), and in Sonneratia it is semi-inferior to superior (Backer
and van Steenis 1951; Duke and Jackes 1987; Tobe et al.
1998), being raised to a superior position by late growth of
the upper portion. Punica includes superior to inferior ovaries.
In Punica protopunica they are superior to semi-inferior (Bal-
four 1888) or semi-inferior (Levin 1980, 2007), and in Punica
granatum they vary from semi-inferior to inferior (Tobe et al.
 Fig. 17 A, B, Tetrataxis salicifolia. A, Raphal face; scale bar p 0.2 mm. B, Antiraphal face; scale bar p 0.25 mm. C, D, Trapa natans. C,
Diagram, longitudinal section of flower showing semi-inferior, bilocular ovary in floral cup, two pendulous ovules (hatched), sectioned circular
coronary disc above the stamens. D, Mature drupe-like fruit, four hornlike sepals, and prominent, indurate projecting apex of the drupe; scale
bar p 10 mm. E–G, Woodfordia fruticosa Kurz. E, Antiraphal face; scale bar p 0.2 mm. F, Raphal face; scale bar p 0.2 mm. G, Evaginated
helical epidermal cell trichomes, antiraphal face; scale bar p 50 mm. (A, B from Lorence 1231; C modified from Ram 1956; D from Grant
16449; E–G from Santapda 14076.)

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228 INTERNATIONAL JOURNAL OF PLANT SCIENCES
1998). In Trapa the ovary is initially semi-inferior and becomes
fully inferior as the fruit matures (Ram 1956). In the tree gen-
era Lagerstroemia and Physocalymma, there are slight ten-
dencies in the superior ovary toward a semi-inferior position;
the ovary base is slightly sunken in the receptacle.
An ovarian stipe occurs in 10 genera: Adenaria, Decodon,
Galpinia, Koehneria, Lafoensia, Lawsonia, some Lythrum,
Pehria, Pleurophora, and Sonneratia. The stipe is obscured by
surrounding nectariferous tissue in Decodon, Galpinia, Lyth-
rum, Pleurophora, and Sonneratia.
Variation in ovary locule number in the family ranges from
one to ∼21 (table 1). Eleven genera consistently have two loc-
ules, and 15 genera have between one and six locules. The
genera formerly placed in three satellite families have the high-
est number of locules. Duabanga is typically four to six loculed
but can have up to nine locules. Punica has three to eight or
nine locules, possibly more, with the lower numbers occurring
in the less modified species, P. protopunica. Sonneratia, with
four to ∼21 locules, usually has 15 or 16 locules (Backer and
van Steenis 1948–1954; Duke and Jackes 1987).
Placentation is basically axile, but the placenta differs in the
extent to which it remains an elongate axis with close asso-
ciation or connection to the apex of the ovary. Also varying
is the degree to which the septal walls are present. Genera
classified in different tribes based on the septal and placental
condition are now found to be members of the same phylo-
genetic clade (e.g., in Lythreae vs. Nesaeeae: Rotala and Hei-
mia, Lythrum and Decodon, Woodfordia and Adenaria; Gra-
ham et al. 2005; Morris 2007) and in one instance are
congeneric (Ammannia and Nesaea; Graham et al. 2011). Dis-
cussion of the placenta separate from the septa better focuses
on the variety of conditions that occur in the ovaries.
The central placental axis may remain elongate as the ovary
matures, or it may become globose by development of en-
larged, fleshy, ovule-bearing lobes that protrude into the loc-
ular spaces. In the early stages of ovary development, the pla-
centa is connected from ovary base to ovary apex, but in
globose forms with growth of the ovary and expansion of the
placenta, the connection is frequently lost above the enlarged
lobes (Joshi and Venkateswarlu 1935–1936). Developmental
studies are needed to verify how often this occurs, but our
observations indicate it is a common condition. Genera that
have an elongated axile placenta that extends at fruit maturity
from the base to or nearly to the apex of the ovary are Cuphea,
Decodon, Heimia, Koehneria, Lagerstroemia, Lawsonia, most
Lythrum, Pleurophora, Tetrataxis, and Woodfordia. In Law-
sonia the distal part of the axis above the ovule-bearing portion
is very slender or possibly is disconnected. The ovule-bearing
portion in Lawsonia is restricted to approximately midlevel of
the placenta and consists of intrusive, slender branched lobes
on which ovules are arranged in tight clusters (fig. 18D).
The septa typically become vestigial or lost above the pla-
cental lobes on the ovary wall and may also be absent below
the fertile lobes even to the base of the ovary. With reduction
of the septa and loss of the distal placental connection, plac-
entation then appears free-basal at fruit maturity. Genera with
free-basal placentation are Adenaria, Ammannia, Capuronia,
Crenea, Didiplis, Diplusodon, Galpinia, Ginoria, Lafoensia,
Lourtella, some Lythrum, Pehria, Pemphis, Physocalymma,
and Rotala. In Duabanga, Punica, and Sonneratia both axile
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and parietal placentation occur in the ovary. Also in Duabanga
and Sonneratia, the placental lobes are enlarged and placen-
tation ultimately appears free-basal. In P. granatum placen-
tation is axile at lower levels and parietal at upper levels in
the ovary, and septa are continuous (Tobe et al. 1998). In
Punica protopunica, with a single whorl of carpels, the septa
are not present or not well developed. Placentation according
to Balfour (1888) is parietal with ovules present on the floor
of the locules and by fruit maturity on the locule walls as well.
Trapa is unique in the Lythraceae for apical placentation. Just
two ovules, one vestigial and one fertile, are produced at the
apex of the semi-inferior to inferior ovary (fig. 17C). Germi-
nation of the single large, starchy seed occurs through the pore
of the stylar canal at the fruit apex.
Anatomical studies have shown all septa are “incomplete,”
that is, free from each other and from the ovary apex (Tobe
et al. 1998). Septa develop to varying degrees both in length
on the ovary wall between the base and apex of the ovary and
in width between the ovary wall and the central axis. In length
they may extend to the ovary apex, but they remain unfused
at the apex. Septal walls reach the ovary apex in 15 genera:
Adenaria, a few Ammannia, Capuronia, Decodon, Duabanga,
Galpinia, Heimia, Koehneria, Lagerstroemia, Lawsonia, most
Lythrum, Punica, Tetrataxis, Trapa, and Woodfordia. Rela-
tively thick septa occur in Decodon, Lagerstroemia, Tetrataxis,
and some Lythrum. In Lagerstroemia only, a secondary septal
extension protrudes into the locule midway between the major
septa at approximately one-fifth or less the distance to the
central axis.
Increasingly reduced septa occur in most Ammannia, Cre-
nea, Didiplis, Diplusodon, Ginoria, Lafoensia, some Lythrum,
Pehria, Pemphis, Physocalymma, and Rotala. In Lafoensia
septa are restricted to the base of the ovary. In Cuphea, Lour-
tella, and Pleurophora, two threadlike vestigial septa tenuously
connect the placenta to the ovary wall (fig. 1D). In Diplusodon,
unique lunate septa extend partially up the ovary wall from
two half placentas to define the two locules.
Ovary position, carpel numbers, and placental and septal
conditions in the Lythraceae vary widely. The diverse mor-
phological conditions in the ovaries and fruits of the Lythra-
ceae call for developmental and anatomical studies to provide
a more detailed comparative account of gynoecial development
and to allow the role of gynoecial changes to be critically
evaluated in the phylogenetic history of the family and the
order Myrtales.
Two of the 28 genera of the Lythraceae produce truly zy-
gomorphic flowers. Ovaries of Cuphea and Pleurophora have
zygomorphic, distinctly unequal-sized carpels in strongly to
lightly bilaterally symmetrical floral tubes. The abaxial (ven-
tral) locule in the bicarpellate ovary is reduced in size and in
number of ovules produced (figs. 1D, 2F; Baum and Leinfellner
1951). In Cuphea in the large subgenus Bracteolatae
S.A.Graham, zygomorphy is emphasized by a dorsal nectar
spur and orientation of the stamens to the ventral side of the
floral tube. There is also a tendency to ventral stamen orien-
tation in the Lythraceae in other genera with tubular flowers
such as Lythrum, Pehria, and Woodfordia. Woodfordia is
slightly zygomorphic, with a slightly smaller ventral locule in
the more distal part of the ovary.
Two genera, Capuronia and Adenaria, are incipiently di-
 Table 1
Summary of Selected Ovary and Fruit Characters of the Lythraceae
Ovary Ovary
Genus Fruit type position stipe Placenta shape Placentation Septa Fruit wall Locules Dehiscence
Adenaria Capsule S Yes Globose FB C Intermediate 2 (3) Irregularly splitting
Ammannia Capsule S No Globose, narrowly ellipsoid FB IC, C Thin (1-) 2–4 (5) Irregularly splitting
Capuronia Berrylike S No Subglobose FB C Thick or intermediate (2-) 3–5 Indehiscent
Crenea Capsule S No Globose FB IC Intermediate 4 (5) Irregularly splitting
Cuphea Capsule S No Ovoid-oblong E V Thin 2 Uniloculicidal
Decodon Capsule S Yes Conical, subglobose E C Intermediate 3 (4) Loculicidal
Didiplis Capsule S No Globose FB IC Thin 2 Irregularly splitting
Diplusodon Capsule S No Bipartite FB IC Thick 2 Septicidal
Duabanga Capsule Semi-I No Subglobose FB, ⫹ parietal C Thick 4–6 (–9) Loculicidal
Galpinia Capsule S Yes Subglobose FB C Thin 2 Irregularly splitting
Ginoria Capsule S No Ovoid, globose FB IC Intermediate 2–6 Septicidal
Heimia Capsule S No Ovoid, subglobose E C Intermediate 4 (5) Loculicidal
Koehneria Capsule S Yes Ovoid, globose E C Thin 2–4 Septicidal
Lafoensia Capsule S Yes Globose to disciform FB IC Thick 2 or 4 Loculicidal
Lagerstroemia Capsule S No Narrowly ovoid E C Thick (3–) 6 Loculicidal
Lawsonia Capsule S No Stipitate, branched E, (FB) C Intermediate (2–) 4 Irregularly splitting
Lourtella Capsule S No Slender ovoid, subglobose FB V Thin 2 Irregularly splitting
Lythrum Capsule S No/yes Slender ovoid, globose E, (FB) C, IC Thin 2 Septicidal
Pehria Capsule S Yes Oblong, globose FB IC Thick 2 Loculicidal
Pemphis Capsule S No Subglobose FB IC Thick 3 or 4 Circumscissile
Physocalymma Capsule S No Subglobose FB IC Thin 4 Irregularly splitting
Pleurophora Capsule S Yes Ovoid E V Thin 2 Irregularly splitting
Punica Berry Semi-II No Twisted or basal-parietal E, ⫹ parietal C Thick 3–8 or 9, more? Indehiscent
Rotala Capsule S No/yes Globose, oblong FB IC Thin 2–5 Septicidal
Sonneratia Berry S, semi-I Yes Subglobose FB, ⫹ parietal IC Thick (4–) 15 or 16 (–21) Indehiscent
Tetrataxis Capsule S No Ovoid E C Intermediate 4 or 5 Septicidal
Trapa Drupe Semi-I, I No Apical apical C Thick 2 Indehiscent
Woodfordia Capsule S No Ellipsoid, narrowly ovoid E C Thin 2 Irregularly splitting
Note. For ovary, S p superior, I p inferior. For stipes, yes p present, prominent; no p not present. For placentation, E p placenta elongate axile, extending to ovary apex at
maturity; FB p placenta appearing free-basal at maturity. For septa, C p septa approximately complete, extending to near ovary apex; IC p septa incomplete, ceasing approximately at
the distal end of the placental lobes or below; V p septa vestigial, forming two thin threads. For fruit walls, thick p more than 10 cell layers; intermediate p moderately thick, 6–10
cell layers; thin p five or fewer cell layers.
 Fig. 18 A, Seeds of selected genera of the Lythraceae illustrating a range in seed sizes and shapes in the family. Lafoensia vandelliana Cham.
& Schltdl, largest seed in the family, forming the background on which smaller seeds of other genera are placed for comparison. The L. vandelliana
seed is discoidal with the central seed body encircled by a broad thin wing; from top to bottom on the Lafoensia seed: Lagerstroemia indica
with obpyramidal seed body and unilateral wing; Cuphea carthagenensis (Jacq.) Macbr., one of several species of the genus having seeds that
are prominently marginate; Decodon verticillatus, obpyramidal, positioned just below middle half of Lafoensia seed; Galpinia transvaalica with

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 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY 231

oecious. In Capuronia, rudimentary ovary or stamens are pres-
ent in the functionally unisexual flowers. In Adenaria, flowers
are dimorphic or weakly trimorphic (Koehne 1903; Nevling
1958). Secretory tissue is described in the Trapa ovary; the
interior ovary wall, placenta, and funicle are “glandular, ex-
uding a gummy secretion” (Ram 1956, p. 313). In Lagerstroe-
mia the epidermis of the placenta is secretory (Odintsova
2008). Globose resin-holding trichomes appear externally on
the distal end of the ovary in the related genera Adenaria,
Koehneria, Pehria, and Woodfordia (fig. 1B; Graham et al.
1986; Graham 1995).
The Fruit
The dominant fruit type in the Lythraceae is a globose to
oblong, pyriform, or obovoid, dry, thin- to thick-walled cap-
sule of two to five (or six) locules that splits irregularly at
maturity. Fruits are partially to fully enclosed in the persistent
perigynous floral cup or floral tube. The thickness of the fruit
wall ranges widely among the genera. Thin-walled fruits are
dry capsules with fragile, brittle, often translucent walls. Ma-
ture thick-walled fruits are leathery and either remain so or
become indurated to heavily indurated and woodlike prior to
dehiscence. Thin-walled fruits occur in Ammannia, Cuphea,
Didiplis, Galpinia, Koehneria, Lourtella, Lythrum, Physoca-
lymma, Pleurophora, Rotala, and Woodfordia (fig. 1D; Cu-
phea). Thick-walled capsules of ∼10 or more layers occur in
Diplusodon, Duabanga, Lafoensia, Lagerstroemia, Pehria,
and Pemphis; in berrylike fruits of Capuronia, Punica, Son-
neratia; and in the drupe-like fruit of Trapa. The pericarps of
Duabanga, Lafoensia, and Lagerstroemia are heavily indu-
rated and woodlike at time of dehiscence. Punica and Son-
neratia fruits are tough and leathery externally and pulpy or
spongy internally. The genera Adenaria, Crenea, Decodon, Gi-
noria, Heimia, Lawsonia, and Tetrataxis have moderately
thickened fruit walls that are typically indurated, tough, or
brittle, not woodlike as in mature fruits of Duabanga, La-
foensia, and Lagerstroemia.
Capuronia fruits are unusual, possibly unique in the Lyth-
raceae. They meet the definition of berries (Lourteig 1985)
being “indehiscent simple fruit[s] with one to many seeds im-
mersed in a fleshy pulp” (Beentje 2012, p. 18). Capuronia fruits
are indehiscent; however, the pericarp cannot be considered
pulpy at all stages of development if observations of fruits from
herbarium specimens are accurate. The pericarp is not differ-
entiated into visibly discernible layers, and there is no hardened
endocarp (fig. 1C). In some fruits taken from exsiccatae, a
thick fruit wall is composed of small compact (mucilaginous?)
cells, and the locules are dry (fig. 1C). In others, the fruit wall
appears thinner, the innermost walls are mucilaginous, and the
locules are filled with a pulp that engulfs the seeds. Mucilage
appears to be released by the breakdown of cells of inner layers
of pericarp. Soaking overnight furthers the breakdown in some
berries. In berries from other exsiccatae, dry mucilage has
“shrink-wrapped” the seed cluster, and it is not reconstituted
by soaking. Studies of fresh material should clarify the true
condition of the fruit. Homology to Sonneratia berries, which
are filled with an acidic pulp, is unknown.
Fruit dehiscence in the Lythraceae is accomplished in several
ways. In 10 of 28 genera the mature fruit splits irregularly,
generally beginning at the fruit apex: Adenaria, Ammannia,
Crenea, Didiplis, Galpinia, Lawsonia, Lourtella, Physoca-
lymma, Pleurophora, and Woodfordia. In Woodfordia dehis-
cence is accomplished by multiple longitudinal splits of the
perigynous floral tube and the included capsule as the floral
tube and capsule enlarge with maturation of the seeds. De-
hiscence is loculicidal in seven genera: Cuphea, Decodon, Dua-
banga, Heimia, Lafoensia, Lagerstroemia, and Pehria. It is
septicidal in six genera: Diplusodon, Ginoria, Koehneria,
Lythrum, Rotala, and Tetrataxis. Indehiscent fruits include
those of Capuronia, Punica, Sonneratia, and Trapa. Uniquely
in Trapa, the single fertile seed in the inferior-ovaried, hardened
fruit ultimately germinates through the neck of the stylar canal
at the apex of the fruit (fig. 17D). Dehiscence is strictly cir-
cumscissile in Pemphis. In many species of Ammannia that
were formerly recognized as Nesaea, irregular splitting is pre-
ceded by an initial circumscissile opening (Graham et al. 2011).
Both thin- and thick-walled fruits in the family split irregularly
from the apex to free a multitude of small seeds. Loculicidal
dehiscence is associated with intermediate- or thick-walled
capsules with the exception of the uniloculicidal, thin-walled
capsules of Cuphea.
The mode of loculicidal dehiscence in Cuphea is a major
diagnostic character of the genus. In the bilocular capsule only
the dorsal locule is dehiscent. It splits along a single dorsal
longitudinal line, and a complimentary longitudinal split oc-
curs on the surrounding persistent floral tube between the two
dorsalmost veins. The capsule remains within the floral tube
while the placenta rises through the slits by means of an in-
crease in turgor pressure and swelling of cells at the base of
the placenta (fig. 18B). Two slender vestigial septal threads
near the placental apex are broken as the placenta bearing
nearly mature green seeds emerges from the floral tube. A
detailed discussion of gynoecial development and the erection
of the placenta in Cuphea is provided by Baum and Leinfellner
(1951). Cuphea seeds require approximately another 2 d fol-
lowing initial exposure to acquire viability, to develop color-
ation in the seed coat, and for the funicular connection to dry.
Seeds
Seed shape is basically obpyramidal or bilateral, although
the shape is not always uniform because packing of seeds in

encircling wing; Ammannia coccinea, lower right, one of the smallest lythraceae seed types; scale bar p 3 mm. B, Dehiscence in Cuphea leptopoda
Hemsl., bottom flower showing initial erection of the elongate placenta through complimentary longitudinal slits in the capsule and floral tube walls,
middle flower with green immature seeds attached (bottom and middle images) on erect placenta, and top flower with fully matured brown seeds,
funicle dried, ready for seed detachment; scale bar p 4 mm (photo by F. Hirsinger). C, Lafoensia punicifolia DC., dry seed on the left showing
central seed body and surrounding flattened wing; water-soaked seed on right with mucilage cells of the wing swollen by absorption of water; scale
bar p 1 cm. D, Lawsonia inermis, fruit partially surrounded by floral cup, longitudinal section through the fruit, placental axis multibranched,
seeds produced in clusters on the branches, septa and extension of the placental axis to the ovary apex not visible; scale bar p 1 mm.

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232 INTERNATIONAL JOURNAL OF PLANT SCIENCES
the capsule or berry alters the shape to varying degrees; ob-
pyramidal forms, especially, can become multifaceted. Bilateral
seeds range from slightly to strongly compressed or with the
antiraphal and raphal faces unequally compressed. Variations
due to position in a tightly filled capsule, shape of the placenta
to which the seeds are attached, moisture state, and other
factors affect details of seed morphology. This should be taken
into account in taxonomic descriptions and particularly in de-
fining new fossil taxa based on seeds. Seeds of Decodon, Lag-
erstroemia (the seed body, not including wing), Lawsonia, and
Pemphis are basically obpyramidal; those of Koehneria vary
from obpyramidal to cuneiform. Bilateral seeds include a range
of seed shapes from nearly globose, obovoid, ellipsoid, or dis-
coidal to fusiform or spindle shaped. In many bilateral seeds,
the antiraphal face is convex whereas the raphal face is flat-
tened to concave, forming a boat shape (e.g., seeds of Adenaria,
Crenea, Lythrum, Rotala, Pehria). In other bilateral seeds, both
sides are convex, being lenticular as in Cuphea, or both sides
are flattened, being discoidal as in Diplusodon, Galpinia, La-
foensia, and Physocalymma. The seed apex (chalazal end) has
a tendency in some bilateral, elongate seeds to become cucul-
late as in Didiplis (fig. 8D, 8E), some Ginoria species (fig.
10A–10C), some Lythrum (fig. 13B), and Tetrataxis (fig. 17A,
17B). Seeds of Sonneratia are basically bilateral but with an
irregular U- or V-shaped outline. A narrow, elongate embryo
and short micropyle occupy one of the two arms of the seed
(fig. 16A, 16B).
Seed numbers range from as few as two per fruit (e.g., in
Cuphea disperma Koehne and Ammannia mossambicensis) to
more than 100 per fruit in many species of Ammannia and
Rotala; in Crenea, Duabanga, Lafoensia, Punica, and Son-
neratia; and in some Cuphea and Lythrum. The large fruits of
Sonneratia and the cultivated Punica and can yield more than
1000 seeds per fruit.
Seeds are minute to small in most genera. Eighteen of 28
genera (64%) produce seeds less than 1 mm long; eight genera
(29%) have seeds with a minimal length of 3.0 mm or greater
(table 2). Small seeds occur in genera of diverse habit and
habitat, in amphibious and aquatic herbs as well as shrubs
and small trees of wet or dry habitats. The largest seeds, ex-
ceeding 4 mm (excluding Trapa), are from tropical trees (La-
foensia, Lagerstroemia, Physocalymma, and Sonneratia).
Wings account for half or more of the seed length in Lafoensia
and Lagerstroemia. The longest seeds are the fragile, slender,
long-tailed seeds of Crenea (5 mm # 0.3–0.5 mm) and Dua-
banga (4–5.5 mm # 0.5 mm). Seed size varies widely in the
species-rich genera Cuphea (0.5–3.3 mm long), Diplusodon
(1–4 mm long), and Lagerstroemia (4.0–16.0 mm long) but
also among the approximately five species of Lafoensia (6.5–
35.0 mm long).
Seed surface patterns are mostly reticulate or scalariform
arrangements of polygonal (tetra- to hexagonal) or rectangular
cells. Surface sculpture is in low relief, provided mostly by the
anticlinal cell walls of the exotesta, which are straight or less
often slightly sinuous, show some difference in thickness
among the genera, and are usually slightly raised in relation
to the typically flat or convex periclinal walls. Periclinal walls
are mostly smooth, infrequently striate when dry. The seed
surfaces in Cuphea subgenus Cuphea, one of the two subgenera
of Cuphea, are more diverse than in any other taxon of the
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family and include the only species with tuberculate and wrin-
kled seed coats (Cuphea circaeoides Smith ex Sims and e.g.,
Cuphea racemosa [L.f.] Spreng. [fig. 6G, 6H], respectively).
The epidermal cells of the Sonneratia seed coat are unusual
in the family. The epidermal layer, composed of thin-walled,
mucilaginous cells, when dry forms a brownish, crusty coat
over a dense spongy mesotestal tissue composed of smaller,
cubical, cream-colored cells (fig. 16C, 16D). When seeds be-
come wet, mucilage in the epidermal cells swells and the cells
separate from the inner spongy layers, falling away as indi-
vidual globose packets of clear mucilage.
Seeds of Decodon are apparently unique for the Lythraceae
in having a prominent germination valve. It is a distinct, more
or less rectangular central area on the antiraphal face of the
seed that becomes detached from the rest of the seed coat at
seed germination. The valve is described from anatomical
transverse sections as attached to the rest of the seed body by
thin connections and is far more prominent on the numerous
fossil seed taxa than on the single modern species (Tiffney
1981). In the rest of the family, a distinct germination valve
that disassociates from the rest of the seed coat has not been
recognized. An area of rectangular cells oriented with the
length of the seed and centered on the antiraphal face over the
seed body is sometimes seen (e.g., Pemphis [fig. 14D], possibly
in Lawsonia [fig. 12A]), but any relationship to a germination
valve has not been investigated anatomically. In several other
genera, longitudinally oriented slightly larger cells occur on
the raphal face in the central raphe region (e.g., Capuronia
[fig. 5B], Didiplis [fig. 8E], some Ginoria [fig. 9F], Lourtella
[fig. 12F]). In Didiplis emergence of the radicle is at the bound-
ary of the two seed faces (fig. 8F). A survey of the anatomy
and germination of Lythraceae seeds is needed to clarify to
what extent the seed coat is modified for germination among
the genera.
Seed Dispersal
Ecologically adaptive structures for dispersal are prominent
features of Lythraceae seeds. Many exhibit morphology indic-
ative of dispersal by water, which is not surprising since the
majority of genera inhabit moist, wet, or fully aquatic habitats.
Buoyancy is achieved by the small seed size, light weight, and
boatlike, discoid, and lenticular shapes. Aquatic dispersal is
assisted in Ammannia (fig. 4F) and Rotala (fig. 15J) by large-
celled aerenchyma on the raphal side of the boat-shaped seeds
and by the extended aerenchymatous tails in Crenea (fig. 5D,
5E) and Duabanga (fig. 7A). The seeds of Adenaria, Decodon,
Lawsonia, Pemphis, Sonneratia, and Tetrataxis are anatomi-
cally adapted for water dispersal by the presence of extensive
internal (mesotestal) float tissue. Detailed descriptions of these
seeds require anatomical studies.
Wind is a primary dispersal agent of winged seeds and is a
typical mode of dispersal for seeds of forest trees where up-
lifting drafts can carry seeds above the tree canopy and away
from the parent plant (van der Pijl 1982). Winged seeds in the
Lythraceae occur in the tropical shrub or tree genera Diplu-
sodon, Galpinia, Lafoensia, Lagerstroemia, and Physoca-
lymma (see fig. 18A for examples from Galpinia, Lafoensia,
and Lagerstroemia). Seeds of the tall tree genera Lafoensia,
Lagerstroemia, and Physocalymma are wind dispersed; they
 Table 2
Summary of Selected Seed Morphological Characters of the Lythraceae

Seeds per Seed length Seed width

Taxon fruit (mm) (mm) Seed shape Wings Trichomes

Adenaria 7–20 .9–1.3 .8–1.2 Bilateral, convex/concave No No

subglobose
Ammannia (2–5) 100⫹ .3–.4 (⫺1.8) .3–.5 (–1.3) Bilateral, convex/concave obovoid- No Straight, ribbed
globose or tubercled
Capuronia 5–8 2.9–3.2 2.3–2.7 Strongly bilateral, cordiform No No
Crenea ∼500 5.0 .3–.5 Bilateral, fusiform to narrowly ob- No Straight, smooth
ovoid, convex/concave
Cuphea 2–100⫹ .5–3.3 .3–3.0 Bilateral, lenticular, ovoid, obo- Rare, encircling, narrow Helical
void, oblong, ellipsoid
Decodon 20–30 1.0–1.9 1.2–2.0 Obpyramidal, 3–6 facets No No
Didiplis 25–45 .6–.9 .2–.3 Bilateral, convex/flattened, elongate no Straight, smooth
obovoid
Diplusodon 4–72 1.0–4.0 1.8–3.5 Bilateral, lenticular to discoidal, Yes, encircling Straight
obovate to orbicular or elliptical
Duabanga 100⫹ 4.0–5.5 .5 Slightly bilateral, fusiform No No
Galpinia 3–10 3.0–4.5 2.5–4 Bilateral, lenticular or discoidal, or- Yes, encircling No
bicular to obovate
Ginoria 50⫹ .5–2.5 .1–.6 Bilateral, convex/concave, oblong, No Straight, smooth
prismatic, obovoid, or fusiform
Heimia 50⫹ .5–1.0 .3–.8 Bilateral, convex/concave, narrowly No Straight, smooth
obovoid-obpyramidal or basal striae
Koehneria 10–30 1.3–1.7 1.0–1.2 Slightly bilateral, obpyramidal to No Helical
approximately cuneiform, 3–4
facets
Lafoensia 100⫹ 6.5–35.0 8.0–15.0 Bilateral, discoidal, subcircular or Yes, encircling Helical
oblong
Lagerstroemia 8–72 3.0–6.0 4.0–16.0 Obpyramidal, wing approximately Yes, unilateral No
rectangular
Lawsonia 30–50 1.2–2.2 1.3–2.0 Obpyramidal, corners rounded No No
Lourtella 30–50 .8–1.0 .5–.7 Obpyramidal, corners rounded No No
Lythrum 30–120 .4–.8 .3–.7 Bilateral, convex/concave or con- No Straight, smooth
vex/flattened, narrowly obovoid, or ribbed
ellipsoid, subglobose
Pehria 50⫹ .7–.9 .3–.5 Bilateral, convex/flattened, nar- No Straight, striated
rowly obovoid by minute
tubercles
Pemphis 15–20 2.5–3.0 1.7–2.0 Obpyramidal, 3–6 facets No No
Physocalymma 50⫹ 3.2–4.2 2.7–4.0 Bilateral, discoidal Yes, encircling No
Pleurophora 20 to ∼100 .6–.7 .4–.5 Bilateral, slightly convex/concave, No Helical
obovoid to ellipsoid
Punica 100⫹ 9.0–12.0 7.0–9.5 Obpyramidal, 3–6 facets No No
Rotala 30–300 .1–1.5 .2–.4 Bilateral, convex/concave, obovoid, No Straight, ribbed,
ellipsoid, subglobose tubercled
Sonneratia 100⫹ to 1300 4.0–5.0 10.0–12.0 Bilateral, flattened or irregularly No No
angular
Tetrataxis 50⫹ .9–1.2 .2–.5 Bilateral, convex/concave obovoid No No
to subcylindric or fusiform
Trapa 1 ∼1.0–2.0a ∼1.0–2.0a Approximately ovoid, retained in No No
fruit
Woodfordia 50⫹ .7–1.0 .2–.3 Slightly bilateral, convex/slightly No Helical
flattened, narrowly obpyramidal
to subcylindric
Note. Seed sizes include wings.
a
In centimeters.

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234 INTERNATIONAL JOURNAL OF PLANT SCIENCES
are lightweight and sturdy with a large surface area. The seeds
of Lagerstroemia, uniquely in the family, possess a unilateral
wing that extends outward along the raphal margin of weighty
obpyramidal seed body, as in samaras. The wing is strength-
ened by a strong top edge of lignified cells (Werker 1997). In
the other genera, the wing is formed by extension of the seed
margin and encircles the bilateral, discoid seed body. In La-
foensia seeds, the epidermis of the seed body consists of short
radially elongated palisade cells whereas the wing is formed
by larger cells with lignified walls (Corner 1976, p. 176). Dis-
coid winged seeds with a central seed body, as in Lafoensia,
Physocalymma, Diplusodon, and Galpinia, in addition to the
lift potential provided by the wing, have increased potential
for aquatic dispersal for the same reasons as for wind dispersal:
their lightweight and enlarged seed surface increase float time
and thus increase dispersal opportunities. Also wind and/or
water dispersed are the fragile fusiform seeds of Crenea, Dua-
banga, Tetrataxis, and some species of Ginoria. All four genera
grow in or near water. Duabanga and Crenea are further cat-
egorized as “dust seeds.” In addition to the small size and light
weight, the slender aerenchymatous tails, extending several
millimeters at each end of the seed, keep the seed longer in
the air (Raub et al. 1975). Duabanga displays the strong an-
ticlinal cell wall pattern seen in many dust seeds (fig. 9B; Barth-
lott 1981).
Several species of Cuphea have narrow encircling seed mar-
gins that have been referred to as “wings” (Koehne 1903).
These seeds are more accurately described as “marginate,” the
encircling margin of the bilateral seed being only slightly ex-
tended and thinned in relation to the rest of the seed coat (figs.
6A, 6B, 7J). The margin appears too narrow to serve as an
effective means of wind dispersal. In aquatic Trapa, the heavy
fruits may initially float and be dispersed by flowing water,
but, ultimately, they sink to the bottom of rivers and ponds,
where they are anchored by the strong curved spines (van der
Pijl 1982); the plants also spread widely by vegetative
fragmentation.
Cuphea seeds offer an opportunity for secondary dispersal
by epizoochory. Because the mature seeds are exposed on the
placenta outside the floral tube and are heavy in relation to
seeds of other genera in their size range, they tend to fall under
or very near the parent plant; namely, they are initially gravity
dispersed (S. A. Graham, personal observation). The embryo,
which fully occupies the internal space of the seed, is high in
oil content (to 42% of seed weight; Graham 1989; average
30% in 30 species analyzed; Graham and Kleiman 1987) and
contains some protein, making them a nutritive wild food re-
source attractive to small birds, rodents, and ants who forage
in ground litter. Evidence that quail ingest Cuphea seeds has
been documented (Hurst 1978). Dry Cuphea seeds were con-
sumed in such numbers by the birds that when the seeds be-
came wet in their crop, mucilaginous trichomes formed large
seed impactions, blocking the digestive passage and leading to
emaciation and probable death. The seed embryos contain sub-
stantial amounts of linoleic acid. Ants are attracted to seeds
in other families of plants that produce oleic and linoleic oil–
rich elaiosomes, carrying seeds to nests up to 10 m from the
parent plant for consumption of the oil-producing organ
(Mayer et al. 2005; Pizo et al. 2005). Although no observations
have been recorded, it would not be surprising to find Cuphea
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seeds ingested by animals or sequestered in ant nests or rodent
middens because of their food value and the ease of access to
the seeds on the ground.
Biotic dispersal is also indicated in Punica by the syndrome
of attractive bright-colored fruits and juicy red seeds. The seeds
are a likely food source for birds and other animals in the wild
(van der Pijl 1982). Capuronia and Sonneratia produce berries
with an acidic pulpy interior, but neither the fruit nor the seeds
are brightly colored. Proboscis monkeys in southeastern Asia
are widely reported to feed on Sonneratia alba leaves, but they
apparently do not eat the fruits. People consume Sonneratia
fruits in Asian dishes for their sour effect (Duke 1983).
Mucilaginous Seeds of the Lythraceae
A specialized feature common in seeds of the Lythraceae is
the ability to produce mucilage. This phenomenon (myxo-
spermy, myxodiaspory) occurs in at least 86 angiosperm fam-
ilies (Grubert 1974, 1981). The mucilage is typically produced
either by trichomes on the surface of the seed coat (e.g., Acan-
thaceae [Daniel and Figueiredo 2009, fig. 10], other taxa [Kapil
et al. 1980]) or under the cuticle layers or as secondary wall
thickenings in the outer epidermal (exotestal) or inner (me-
sotestal) cells in many taxa (e.g., Brassicaceae; Vaughan and
Whitehouse 1971; Western et al. 2000).
In the Lythraceae, uniquely among all angiosperms, muci-
lage is produced by a single straight or helical trichome that
develops to the inside of the outer wall of the epidermal cells
of the seed coat (Stubbs and Slabas 1982). Upon wetting of
the seed, the trichome absorbs water, swells, a circular lid in
the cell wall over the attachment area ruptures, and the tri-
chome evaginates, releasing expanding mucilage as it elon-
gates. Koehne (1885, p. 33), with reference to an earlier report
on Lythrum and Peplis seed trichomes by Kiärskou (1874),
recorded the presence of the swelling seed trichomes in Am-
mannia, Crenea, Cuphea, Lythrum, Nesaea, and Peplis. Cor-
rens (1892) and Gruetter (1893) first described the develop-
ment of the trichomes. Since then, little attention has been paid
to the trichome character in the Lythraceae. This study is the
first to survey all genera of the family for this phenomenon.
Fourteen of the 28 genera of the Lythraceae form epidermal
internal mucilaginous trichomes. The trichomes are straight in
nine genera (Ammannia, Crenea, Didiplis, Diplusodon [T.
Cavalcanti, personal communication], Ginoria, Heimia, Lyth-
rum, Pehria, and Rotala) and helical in five (Cuphea,
Koehneria, Lafoensia, Pleurophora, and Woodfordia; table 2).
The majority of seeds less than 2 mm long have mucilaginous
trichomes (table 2). Stubbs and Slabas (1982) found that the
helical trichomes of Cuphea procumbens Ortega consisted of
a proteinaceous wall that probably maintains the helical struc-
ture and a polysaccharide core mainly composed of arabinose,
ribose, and xylose. The polysaccharides are produced by Golgi
bodies in the epidermal cells and are secreted to the cell wall
or can remain in the cell space, as happens in some cells of
Lafoensia. Trichome lengths vary among the genera from ∼50
mm to 2 mm. (Additional information on seed trichome lengths
in species of the larger genera is needed for a more accurate
estimate of variation.) Straight trichomes in small seeds are
typically less than 200 mm long; they are 800–1200 mm in the
relatively large seeds of Diplusodon. Helical trichomes are typ-
 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY
ically longer than 200 mm in the helical form and longest in
seeds of some Cuphea (to 2 mm; fig. 7E) and Lafoensia (600
mm; fig. 11F).
The roles played by seed mucilage in seed dispersal, viability,
and germination have been widely reviewed, and in recent
years new roles have been proposed, including DNA repair in
the seed embryo (Grubert 1974; Ryding 2001; Sun et al. 2012;
Western 2012; Yang et al. 2011, 2012). However, controlled
studies to precisely delineate the ecological roles of plant mu-
cilage remain limited (Western 2012). Some studies have been
directed to the numerous species with mucilage seed that occur
in arid environments. Seeds of Plantago (Gutterman and Shem-
Tov 1996; Wu et al. 2009), Alyssum (Sun et al. 2012), and
Artemisia (Yang et al. 2010), among others, under osmotic
stress in arid conditions germinate faster and with higher rates
of germination in the presence of seed mucilage than without
mucilage. Of the seven genera of the Lythraceae primarily or
exclusively inhabiting semiarid shrublands, three produce mu-
cilaginous trichomes (Diplusodon, Koehneria, and Lafoensia)
and four do not (Capuronia, Lawsonia, Lourtella, and Pun-
ica). Any evolutionary advantage of the structurally more com-
plex helical trichomes that appear in two of the most derived
clades of the family (see phylogenetic considerations) is not
clear; one possibility is that greater lengths of the helical form
can be packed within a cell, thus favoring seed survival by the
increased amount of mucilage stored and thus increased ca-
pacity for water absorption.
Drying mucilage cements seeds to surrounding soil particles
increasing soil/seed mass stability and thus decreasing distri-
bution of seeds to unfavorable habitats or dispersal by wind.
If seeds become affixed to animal fur, feathers, or feet by mu-
cilage, they could be widely dispersed (Gutterman 1994; West-
ern 2012; Yang et al. 2012). Such attachment is sometimes
cited as a means of long-distance dispersal; however, we know
of no published observations to that effect for seeds of the
Lythraceae. It is likely that seed mucilage in this family typically
clumps and fixes seeds to moist habitats. Germination is then
enhanced by increased water uptake, which softens or breaks
the hardened internal sclerotic and lignified cell layers of the
seed that characteristically occur in the Lythraceae, allowing
easier release of the radical from the seed coat. The fully evag-
inated mucilaginous seed trichomes of Lythraceae seeds may
also aid aquatic dispersal by expanding the float surface area.
Trees with winged, wind-dispersed seeds might not be ex-
pected to expend energy on mucilage production if long-dis-
tance dispersal by wind was the only adaptive influence on
seed form. That is the case in three of the five large tree genera,
Duabanga, Lagerstroemia, and Physocalymma, that have
winged seeds and lack mucilage trichomes. Seeds of Lafoensia,
however, have wings and produce mucilage in both helical
trichomes and in epidermal mucilage cells. The dual adapta-
tions enhance opportunities for seed dispersal and for in-
creased water absorption needed to ensure seed germination
and establishment in the open woodlands, grasslands, and dry
cerrado favored by this genus. Seeds of the fifth tree genus,
the mangrove Sonneratia, are not winged, produce mucilage,
and are water dispersed. The adaptive significance of mucilage
in this genus is elusive. A summary of ovary and fruit char-
acters of the Lythraceae is provided in table 1 and characters
of seed morphology in table 2.
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235
Phylogenetic Characterization
The focus of this study is to describe the morphological
diversity of poorly known or previously undescribed charac-
ters in ovaries, fruits, and seeds of the Lythraceae. A compre-
hensive treatment of the evolution of these characters is re-
served for future phylogenetic investigation when associated
anatomical characters can be included. However, here we
briefly address the evolution of six morphological characters
of likely significant adaptive value in the spread and diversifi-
cation of the family. Character states for ovary position, pla-
centa type, locule size, fruit types, seed wings, and seed tri-
chomes (tables 3, 4) were parsimoniously optimized on a
phylogenetic tree projected from earlier analyses (see Meth-
odology; Maddison and Maddison 1992). The results are dis-
cussed, and four characters mapped to the tree are illustrated
(fig. 19A–19D).
Ovary position. Until recent phylogenies informed rela-
tionships, ovary position distinguished the superior-ovaried
Lythraceae from the similar but inferior- or semi-inferior-ova-
ried Duabangaceae, Onagraceae, Punicaceae, Sonneratiaceae,
and Trapaceae (Dahlgren and Thorne 1984). The first mor-
phological phylogenies produced for the Myrtales (Johnson
and Briggs 1984) followed the unidirectional, superior to in-
ferior evolutionary trend in ovary position standardly accepted
in angiosperm evolution. The superior ovary occurred in the
ancestor common to Lythraceae, Duabangaceae, Punicaceae,
and Sonneratiaceae. It was retained in the Lythraceae sensu
stricto but evolved toward inferiority in Duabanga, Punica,
and Sonneratia. The semi-inferior- to inferior-ovaried Trapa-
ceae and Onagraceae diverged as a separate branch from Lyth-
raceae-Duabanga-Punica-Sonneratia. Although the trend from
superior to inferior remains the common evolutionary direc-
tion, the evolution of ovary position is now understood as
more dynamic and diverse (Soltis et al. 2003). Based on more
recent phylogenetic analyses of the Myrtales (Sytsma et al.
2004), the ovary position of the ancestor to the Lythraceae is
equivocal. The three major lineages of the family forming an
unresolved tritomy at the base of the family (Graham et al.
2005) are plesiomorphic for superior ovaries, and variants
from the superior position are confined to members of lineage
II, clade 3 (Duabanga, Sonneratia, Trapa), and lineage III,
clade 5 (Punica; fig. 19). The four genera are strongly sup-
ported as members of a monophyletic Lythraceae based on
multiple molecular evidence (Graham et al. 2005). Some of
the variation leading to a fully inferior ovary in Punica gran-
atum was probably influenced by selection of cultivars for
larger, more protected, more productive fruits in that species.
Placenta. The placenta is one of the most highly diversified
floral morphological features in the Lythraceae. The plesio-
morphic condition, occurring in 11 genera, is an elongate axis
that remains elongate as the ovary reaches maturity. It is con-
nected by septa to the ovary wall over all or most of its length.
In the rest of the family, the initial continuous attachment of
the placenta from base to ovary apex is lost by seed maturity.
Enlargement of placental lobes occurs on both the elongate
placental axis and the abbreviated free-basal form, although
the lobes appear to protrude less into the locules on the elon-
gate axis. Free-basal placentation has evolved in all three lin-
eages in four of six clades. In lineage I it arose once. In lineage
236 INTERNATIONAL JOURNAL OF PLANT SCIENCES

Table 3 The morphological diversity in seeds among taxa of a single

clade, even between sister genera, is striking. Examples of di-
Coding for Character Analysis
vergence in seed types held by sister genera can be seen in
Character Character coding Decodon and Lythrum, Duabanga and Lagerstroemia, La-
1. Ovary position 0 p superior; 1 p semi-inferior; foensia and Punica, and Lourtella and Physocalymma. The
2 p inferior distinctiveness in seeds of closely related taxa extends also to
2. Ovary locules 0 p equal in size; 1 p unequal in size the seed anatomy and is a separate subject of investigation.
3. Fruit type 0 p capsule; 1 p berrylike; Ovary locules and fruit types. Changes were also mapped
2 p drupaceous on the family phylogeny to trace the evolution of unequal
4. Placenta 0 p elongate axile at maturity; locule development and diverse fruit types (not illustrated).
1 p free basal axile at maturity; The reduction in the ventral locule of a bilocular ovary has
2 p apical; 3p parietal
occurred only in lineage III, clade 6; once in the immediate
5. Seed wings 0 p absent; 1 p encircling;
2 p lateral
ancestor of South American sister genera Pleurophora and Cu-
6. Seed internal trichomes 0 p absent; 1 p straight; 2 p helical phea; and independently but less strongly expressed in the
Afro-Asian Woodfordia. The flowers of these genera share a
tubular form and bilaterally symmetry, with stamens emerging
II it arose independently twice. Half (five) of the genera retain or tending to emerge on the ventral side of the tube. The
elongate placentas whereas in the other five (clades 3 and 4) smaller ventral locule may allow the proboscis of a hovering
diversification includes free-basal, apical, and mixed axile/pa- insect visitor more open access to nectar at the base of the
rietal forms. In lineage III, change to free-basal placentation floral tube. At the same time, versatile dorsally oriented anthers
evolved a minimum of four times: once in clade 5 and three project from the ventral side of the tube, in position for transfer
times in clade 6. The change appears in both Old World and of pollen to the underside of a visiting insect. Woodfordia
New World genera. The apical production of ovules in Trapa shows less bilateral symmetry of the floral tube and less re-
is unique in the Lythraceae; it is also diagnostic of a member duction in the ventral locule than the other genera. The sister
of the outgroup, Combretaceae (Stace 2007). The morpho-
logical diversity in placental form and the degree to which the
septa are present indicate the need for developmental and an- Table 4
atomical studies to more clearly understand the variability and Morphological Data Matrix
change that occurs during maturation of the ovary.
Character
Seed wings. Nonwinged seeds are plesiomorphic, occur-
ring in 21 genera (including Crenea, of unknown phylogenetic Taxon 1 2 3 4 5 6
position). Unilateral-winged seeds evolved only once, in Lag- Adenaria 0 0 0 1 0 0
erstroemia in lineage II. Encircling wings, restricted to four Ammannia 0 0 0 1 0 1
genera of lineage III, are derived four times independently from Capuronia 0 0 1 1 0 0
the nonwinged state in clades 5 and 6. Diplusodon, Lafoensia, Crenea 0 0 0 1 0 1
and Physocalymma are trees and shrubs of dry or seasonally Cuphea 0 1 0 0 0 2
dry cerrados and savannas, and Galpinia is a small tree of Decodon 0 0 0 0 0 0
forest margins where seed dispersal by wind is to be expected. Didiplis 0 0 0 1 0 1
Alternatively, in clade 5, seeds of the nonwinged genus Pem- Diplusodon 0 0 0 1 1 1
Duabanga 1 0 0 0,3 0 0
phis, an inhabitant of ocean coasts, were adapted for aquatic
Galpinia 0 0 0 1 1 0
dispersal by procurement of internal float tissue, and Punica Ginoria 0 0 0 1 0 1
and Capuronia, genera of dry vegetation types, became Heimia 0 0 0 0 0 1
adapted for epizooic dispersal in the gain of berry-type fruits. Koehneria 0 0 0 0 0 2
In clade 6 where seven of the nine genera lack wings, six are Lafoensia 0 0 0 1 1 2
provided with mucilaginous trichomes. Lagerstroemia 0 0 0 0 2 0
Seed trichomes. Mucilaginous inverted seed trichomes ap- Lawsonia 0 0 0 0 0 0
pear in all lineages and clades except clade 3. The simple, Lourtella 0 0 0 1 0 0
straight trichomes are a synapomorphy for lineage I; straight Lythrum 0 0 0 0 0 1
trichomes also have evolved in the other two lineages. The Pehria 0 0 0 1 0 1
Pemphis 0 0 0 1 0 0
helical trichome type is restricted to lineage III, in clade 5 in
Physocalymma 0 0 0 1 1 0
Lafoensia and in clade 6 where it is independently derived in Pleurophora 0 1 0 0 0 2
four genera, either once or twice from an ancestor lacking seed Punica 0,1,2 0 1 0,3 0 0
trichomes, and in the Old World and the New World. An Rotala 0 0 0 1 0 1
independent gain of straight seed trichomes also occurs in clade Sonneratia 0,1 0 1 0,3 0 0
6 in Adenaria. Diplusodon, like Lafoensia, has seeds with both Tetrataxis 0 0 0 0 0 0
encircling wings and trichomes (T. Cavalcanti, personal com- Trapa 1 0 2 2 0 0
munication). Rotala and Ammannia have independently Woodfordia 0 1 0 0 0 2
evolved both inverted trichomes and large-celled epidermal Outgroup-Ludwigia 2 0 0 0 0 0
aerenchyma. Crenea, absent from the phylogenetic tree, also Outgroup-Fuchsia 2 0 0 0 0 0
Outgroup-Combretum 2 0 2 2 2 0
has trichomes and epidermal aerenchyma.

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 Fig. 19 Reconstruction of four selected morphological characters on a multigene-based phylogeny of the Lythraceae (Crenea absent). Three
lineages are unresolved at the base of the family; six clades are numbered. Genera of Old World origin are shown in bold; those of New World
origin are in roman type; gray striped branches are equivocal. A, Ovary position. B, Placenta type. C, Seed wings. D, Seed trichomes.

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238 INTERNATIONAL JOURNAL OF PLANT SCIENCES
genus to Woodfordia, Koehneria, has a floral cup and is ra-
dially symmetrical with equal-sized locules.
The evolution of fruit types from the plesiomorphic capsular
form is limited to berrylike fruits and the unique drupe-like
fruit of Trapa. Berry types appear in three genera in two lin-
eages (Sonneratia, lineage II, clade 3; Punica and Capuronia,
lineage III, clade 5). The morphologies of the three berrylike
fruits are sufficiently dissimilar to suggest they are not
homologous.
From the phylogenetic viewpoint, the diverse ovary, fruit,
and seeds examined either are autapomorphies on terminal
branches or have evolved repeatedly across the family. Ho-
moplasy is the rule. None of the characters investigated in this
study is a unique synapomorph for any of the three major
lineages or clades. Little diversification is apparent on most of
the lower branches of the phylogeny; the majority of changes
map to terminal nodes. Changes to free-basal placentation,
production of encircling seed wings, and inverted straight and
helical seed trichomes have arisen on multiple occasions in
both the Old World and the New World. Evolution of semi-
inferior and inferior ovaries from superior ovaries and the rare
occurrences of fruit types other than capsules are restricted to
genera of the Old World. Among the clades, clade 5 is excep-
tionally diverse. Genera, four endemic to the Old World and
one to the New World, include superior and inferior ovaries;
capsular and berrylike fruits; fruit dehiscence loculicidal, cir-
cumscissile, by irregular splitting, or indehiscent; and seeds
with or without wings or with spongy float tissue or a sar-
cotesta. Clade 3, of Old World genera, is most diverse with
respect to ovary position and placentation type.
The Historic Fruit and Seed Record
The Lythraceae have a relatively extensive fossil record that
includes vegetative organs, flowers, pollen, fruits, and seeds.
Fossils range in age from 82 million Ma to the Holocene (Gra-
ham 2013). Twelve genera of the family are represented by
fossil fruits and/or seeds; five are extant (aff. Adenaria, De-
codon, Lawsonia, Punica, and Trapa), and seven are extinct
(Alatospermum Chandler, Hemitrapa Miki, Microdiptera
Chandler, Minsterocarpum Reid & Chandler, Pachyspermum
Reid & Chandler, Enigmocarpon Sahni, and Shirleya Pigg &
DeVore; Graham 2013).
The oldest fossils of unquestioned Lythraceae ancestry are
pollen grains of Lythrum/Peplis from the Late Cretaceous early
Campanian of Wyoming (82–81 Ma; Grimsson et al. 2011;
lineage II). Modern pollen of Lythrum is nearly identical to
fossil Lythrum pollen, and habitats are now, as they were then,
wetlands of various kinds, allowing inference that if modern
and fossil pollen are identical, modern and fossil Lythrum
seeds may also be similar. That is, earliest Lythrum seeds would
have been very small, basically bilateral, obovoid, with straight
mucilaginous trichomes, and water or possibly bird dispersed.
In the same line of reasoning, inverted seed trichomes can be
inferred in the fossils of four other modern trichome-bearing
genera with fossil pollen or leaf records, Crenea (fossil pollen,
late Eocene), Cuphea (fossil pollen, early Miocene), Lafoensia
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(pollen, Holocene), and Woodfordia (leaves, Pleistocene). This
suggests that inverted straight seed trichomes, as in Lythrum
and Crenea, were an early and persistent seed innovation in
the family and the helical form in four members of clade 6
and Lafoensia of clade 5 are a later innovation dating to as
early as the early Miocene.
Next-oldest fossils of the Lythraceae are seeds of Decodon
from the late Campanian of northern Mexico (73.5 Ma;
Estrada-Ruiz et al. 2009). Decodon has one of the most ex-
tensive fossil records in the family, and the record is primarily
based on fossil seeds (Graham 2013). Fossil Decodon seeds
also occur in North America in the middle Eocene of British
Columbia (∼45–50 Ma; Cevallos-Ferriz and Stockey 1988)
and are common and most diverse in Miocene and Pliocene
strata of Europe and Asia (Grimsson et al. 2011). Spongy float
tissue is well developed in fossil Decodon seeds and in four
similar extinct seed genera from the Eocene (Alatospermum,
Microdiptera, Minsterocarpum, and Pachyspermum; Graham
2013) and persists in modern Decodon. Fossil Sonneratia
(wood, early Paleocene, ∼67 Ma) by inference had bilateral
irregularly shaped, multitudinous, water-dispersed seeds, and
Lagerstroemia (leaves, late Paleocene/Eocene, ∼56 Ma) had
wind-dispersed obpyramidal unilateral winged seeds by the
beginning of the Eocene (Graham 2013). Fossil winged seeds
of Shirleya Pigg & DeVore very similar to those of Lagerstroe-
mia are well preserved in the middle Miocene of Washington
State (Pigg and DeVore 2005).
Documentation of the diverse forms of ovaries, fruits, and
seeds in the Lythraceae is significant for three principal reasons:
(1) they provide many important diagnostic characters em-
ployed for taxonomic identification of both modern and fossil
plant specimens, (2) accurate identification of the fossils fa-
cilitates the phylogenetic dating of lythracean lineages by pro-
viding reliable constraint points for divergences, and (3) these
identifications are essential for eventually establishing the time
and place of origin for the lineages and the direction and rate
of migration and in explaining current distributions and geo-
graphic affinities evident in the family. The innovative adap-
tations of the seeds for water, wind, and possible epizoochoric
dispersal were likely already present early in both the New
World and the Old World. Their presence and the 82 million
years or more available to accomplish dispersal played major
roles in the diversification and extensive distribution of modern
Lythraceae.
Acknowledgments
We gratefully acknowledge Olga Martha Montiel and Jack
Regalado for Duabanga seeds; Taciana Cavalcanti for infor-
mation and material of Diplusodon; and Fred Keusenkothen,
Monica Carlsen, Rosa Ortiz, and Alba Arbelaez for valued
technical assistance. Hiroshi Tobe generously permitted use of
anatomical photographs from his unpublished research. The
authors thank the reviewers and the editor for their meticulous
reviews of the manuscript and for helpful, informative
suggestions.
 GRAHAM & GRAHAM—LYTHRACEAE OVARY, FRUIT, SEED MORPHOLOGY
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