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The genus Tubocapsicum (Solanaceae)

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Tubocapsicum (Solanaceae: Solanoideae), a genus of two Asian species, was formerly included in Capsicum. Tubocapsicum anomalum is a herb with small yellow flowers and red berries that is found in Japan, southern China, Taiwan, and the Philippines; T. obtusum is found in Japan. A general taxonomic description is complemented by new detailed descriptions and illustrations of growth morphology, vegetative histology and stomata, floral structure, vasculature, fruit structure, seeds, and pollen. On the basis of this analysis as well as cpDNA investigations by of Olmstead et al. (1999), the genus Tubocapsicum appears to be most closely related to subfamily Solanoideae genera Aureliana and Withania.
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The genus Tubocapsicum (Solanaceae)
William G. D’Arcy1, Richard C. Keating1, Zhi-Yun Zhang2, and Ching-I Peng3,*
1Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166, USA
(William G. D’Arcy passed away on December 16, 1999)
2 Institute of Botany, Chinese Academy of Sciences, Xiangshan, Nanxincun 20, Beijing 100093, China
3Institute of Botany, Academia Sinica, Taipei 115, Taiwan
(Received November 23, 1999; Accepted June 22, 2000)
Abstract. Tubocapsicum (Solanaceae: Solanoideae), a genus of two Asian species, was formerly included in Capsicum.
Tubocapsicum anomalum is a herb with small yellow flowers and red berries that is found in Japan, southern China,
Taiwan, and the Philippines; T. obtusum is found in Japan. A general taxonomic description is complemented by
new detailed descriptions and illustrations of growth morphology, vegetative histology and stomata, floral structure,
vasculature, fruit structure, seeds, and pollen. On the basis of this analysis as well as cpDNA investigations by of
Olmstead et al. (1999), the genus Tubocapsicum appears to be most closely related to subfamily Solanoideae genera
Aureliana and Withania.
Keywords: Anatomy; Anther; Branching; Capsicum; Floral vasculature; Fruit; Growth pattern; Nectar; Pollen;
Revision; Seeds; Solanaceae; Stomata; Taxonomy; Tubocapsicum.
Introduction
The genus Tubocapsicum Makino is confined to east-
ern Asia (Figures 1, 2). It embraces one widespread
species, T. anomalum (Franch & Sav.) Makino and a
poorly known second species, T. obtusum (Makino)
Kitamura, of Japan. The genus is not well known since
there has been little published information, and there are
few specimens in western herbaria. In order to redress
this obscurity, we undertook a study of living plants of
the genus cultivated from seeds from Taiwan, as well as
herbarium material from throughout its range. Here we
report our observations and provide a treatment of the ge-
nus Tubocapsicum with descriptions, drawings, photos,
and an analysis of its possible relationships within the
Solanaceae. Nearly all of the observations on the struc-
ture and biology of the genus to follow are of T. anomalum
as T. obtusum is only known thus far from a few herbarium
specimens.
Plants of Tubocapsicum anomalum (Figures 3, 4, 5, 6)
are erect or sprawling herbs with entire leaves, few-flow-
ered inflorescences, small campanulate flowers, and red
juicy berries with discoid seeds. In most features, they
resemble members of Acnistus Schott, Aureliana Sendt.,
Capsicum L., Vassobia Rusby, Witheringia L’Her. and
other genera of the American tribe Capsiceae. However,
as discussed below, seed comparisons and cpDNA stud-
ies (Olmstead et al., 1999) argue for separating
Tubocapscium and Aureliana from tribe Capsiceae, along
with Withania Pauq. which has been placed in tribe
Physaleae.
In this paper we present a summary of our observations
of living plants and herbarium material of Tubocapsicum
anomalum. We include a report on germination and
growth patterns, vegetative histology and vasculature, flo-
ral structure and vasculature, fruit structure, pollen, seeds,
quality of the nectar, notes on its cultivation, and a gen-
eral discussion of how it differs from other genera. We
conclude with a systematic revision based on literature and
herbarium collections including nomenclature of the genus,
its two species, and a map of its distribution.
Bot. Bull. Acad. Sin. (2001) 42: 67-84
90E 100E 110E 120E 130E 140E
30N
25N
20N
15N
10N
5N
35N
40N
Figure 1. Distribution of Tubocapsicum anomalum. Solid
squares indicate specimens seen by the authors. Large circles
indicate reports from the literature.*Corresponding author. E-mail: bopeng@gate.sinica.edu.tw
68 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
Figure 4. Tubocapsicum anomalum (Franch. & Sav.) Makino. A, Inflorescence with flower buds; B, Inflorescence with flower in
lateral view; C, Flower in frontal view; D, Fruit.
Figure 3. Mature plant of Tubocapsicum anomalum (Franch.
& Sav.) Makino reclining to the horizontal.
Figure 2. Distribution and altitudinal maps of Tubocapsicum
anomalum (dots) in Taiwan.
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 69
Materials and Methods
Plants of Tubocapsicum anomalum grown from seed
from Taiwan (Chen 231, HAST, MO) in greenhouses and
outdoors at the Missouri Botanical Garden, St. Louis,
Missouri, USA, for several years were studied morpho-
logically and anatomically and compared with plants of
its putative relatives, also grown there. All cultivated
specimens are vouchered and deposited at MO.
Comparative Living Material Studied: Capsicum
annuum L. var. annuum. Various commercial seeds and
fruits available in markets in St. Louis. not vouchered;
Capsicum annuum var. aviculare (Dierb.) D’Arcy &
Eshb., Mexico, Sonora, La Argentina, seed from W. H.
Eshbaugh, D’Arcy 17723 (MO); Capsicum lanceolatum
(Greenm.) C.V. Morton & Standl., Mexico, Capes, Tay-
Figure 6. Tubocapsicum anomalum. A, Habit; B, Flower; C, Flower from the front; D, Flower bud; E, Opened flower showing
pistil and part of androecium; F, Stamen, ventral (adaxial) view; G, Stamen, dorsal (abaxial) view; H, Fruit.
Figure 5. Interior of Tubocapsicum anomalum flower show-
ing anthers in erect position.
70 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
lor & Breelove 41082 (MO); Tubocapsicum anomalum,
seeds from Taiwan, Nantou Hsien: Fenghuangshan, 1280
m, 24 Oct 1987, Chen 231 (HAST), vouchered as D’Arcy
19501 (MO); Vassobia breviflora (Sendt.) Hunz.,
Argentina, Prov. Córdoba, D’Arcy & Hunziker 13949
(MO); Withania riebeckii Balf. f., Socotra, seed from
Conservatoire et Jardin Botanique de Nancy, D’Arcy
17750 (MO); Withania somnifera (L.) Dunal, source
unknown, D’Arcy 17745 (MO).
A comprehensive sample of herbarium specimens of
Tubocapsicum and putative relatives were also studied as
cited below. Except specimens used to judge the validity
of T. obtusum, all material studied was of T. anomalum
var. anomalum.
Seeds of Tubocapsicum, Vassobia, and Withania were
prepared for study by a 30-second extraction from fresh
berries using a juice blender, followed by air drying. Seeds
of Aureliana were prepared in the same way from intact,
unflattened berries taken from herbarium sheets and
rehydrated. Hand sections of seeds and leaves were
stained in 15% ethanolic iodine-potassium iodide (2.0%
potassium iodide and 0.2% iodine) and mounted in 40%
calcium chloride, which is a stain-differentiating mountant
with a high refractive index (Herr, 1992; Keating, 1996).
Nectar sugar ratios were obtained by C.E. Freeman
(pers. comm.) using the protocol outlined in Freeman et
al. (1984).
Observations
Germination
Tubocapsicum anomalum is phanerocotylar, the seed-
ling often emerging completely from the testa. The coty-
ledons are ovate, with slight differentiation of the distal
region into an area suggestive of an apicule. The cells of
the upper epidermis are quadrate, uniform, and oriented
perpendicular to the midvein. Cells of the lower epider-
mis are rounded and less regularly oriented, with those
near the midvein arranged parallel to it. Cotyledon ve-
nation is irregular, looped, brochidodromous. Secondary
veins are irregularly and widely spaced. There are sparse
uniseriate trichomes on both surfaces. The petiole is
slender, slightly flattened, about 3/4 as long as the blade.
The stem and petioles of the cotyledon are beset with
minute, transparent unicellular subconical trichomes.
The eophylls are alternate, petiolate, estipulate, linear
or oblong, the lateral veins ca. 2 on each side of the
midvein, ascending, the upper (adaxial) surface, petiole
and stalk with tiny, linear, transparent unicellular hairs
that sometimes bear apical glands. These details conform
to those reported by Duke (1969) for the Solanaceae, and
they also correspond with our observations of seedlings
of Capsicum lanceolatum, Vassobia breviflora, Withania
somnifera, and W. riebeckii, except that in these the
eophylls are much broader.
Adult Growth Architecture
After germination, T. anomalum plants develop though
branching or structural phases (Figure 7) as follows. Fol-
lowing development of the eophylls, the plant develops a
straight, relatively stout, erect trunk with a large pith. In
this orthotropic or monopodial phase, leaves are inserted
in a spiral 2/5 phyllotaxy, and axillary buds or shoots are
usually not evident. After several leaf nodes, at a height
of about 30-50 cm, an inflorescence is produced, subtended
on each side by a plagiotropic (sympodial) branch, and
these branches are subtended by a single leaf. Each
Figure 7. Adult growth patterns of Tubocapsicum anomalum. (A) Early maturity: Growth begins with an orthotropic phase which
results in a relatively stout, erect trunk with a large pith (1). After appearance of the first inflorescence (2), more or less equal
pairs or triads of plagiotropic branches continue the plant’s growth, each terminating in an inflorescence and another tier of plagio-
tropic branches. The plant is usually erect. (B) Advanced maturity: Distal branches become, thin, weak, and senescent (3). When
a branch declines to the horizontal, the branches it produces are unequal, the lower branch of the pair becomes dominant (4) and
the upper one is reduced in size or subordinate to it (5). Reiteration shoots (6) that resemble the initial orthotropic trunk appear
from near the base of the plant. SE = senescent branches, SU = subordinate branches.
B
A
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 71
branch in turn grows on to produce an inflorescence sub-
tended by a tier of branches. Under favorable growing
conditions, each subsequent node usually displays this
2-tiered or dichasial arrangement or rarely a 3-tiered
arrangement, though sometimes only a single subtending
branch appears. The two or three plagiotropic branches
appear to be parallel in development, neither showing
dominance. These branches in turn end in inflorescences
and tiers of branching, and they tend to have only occa-
sional leaves except at the branch nodes. The branching
pattern here is dichasial and the sympodial unit is unifoli-
ate (Knapp, 1989: 66).
After a succession of 2-5 branch nodes, the branches
are often thinner, and senescence may appear in the dis-
tal 1-2 nodes of one of the branches of a node. At about
this time, reiteration or rejuvenation shoots appear, usu-
ally at the base of the plant. These shoots resemble the
initial, stout, orthotropic trunk except that they have much
larger leaves than the older shoots, and they subtend ax-
illary shoot initials near the base of the parent trunk.
These shoot initials are evident as fascicles of 2-6 small
leaves that do not develop further. The first dichotomy
of the reiteration shoot occurs after 6-13 leaf-nodes and
is terminated by an inflorescence as in the initial
orthotropic trunk.
Branching architecture of the plant tends to be plastic,
and a number of modifications to the above pattern are
found. Commonly, after the initial period of growth just
described, the stems of Tubocapsicum anomalum are weak
and spindly, and originally erect branches may decline
in part or entirely to the horizontal or arch downwards.
When a branch is no longer close to erect, the branch of
the pair that is lowermost (closest to the ground) tends to
grow faster and it soon assumes dominance in the pair.
The more erect, subordinate, branch of the pair may cease
elongation and remain as a short-shoot or fascicle of
leaves, while the dominant, more horizontal member
arches upward, its base horizontal and its apex assuming
a more vertical orientation. The dominant lower branch
often does not attain a fully erect position, and its subse-
quent nodes are also somewhat horizontal with their
branch pairs, in turn, also showing inequality and domi-
nance by the lower member. Thus, a lengthy but spindly
trunk may be formed with leafy fascicles at each node rep-
resenting the shorter members of tier pairs.
Another common growth form modification occurs
when branches decline greatly from the vertical, the emer-
gence of one or more reiteration shoots from the axils of
leaves on the primary or parent stem. If the plagiotropic
parent branch arches downward, these reiteration shoots
tend to form on the highest part of the arch. Because
plants are usually unable to maintain erect growth after
the first few tiers of plagiotropic branches are produced,
plants that are past their initial flowering period of 2-5
dichotomies tend to have a mixture of senescing plagio-
tropic shoots and reiteration shoots that may have
reinitiated platiotropic branching. At this point there may
be no main trunk, although a number of minor trunks are
present as a result of the dominance of members of tier
pairs. The plant now consists of a number of distinct sym-
podial modules, and the architecture is reminiscent of the
Prevost and Nozeran models (Halle et al., 1978) in which
new modules appear below the first branch tier of the pre-
vious module. In this case, however, the new module may
appear at various places, depending on the vertical ori-
entation of the plant and perhaps other factors, although
under good growing conditions when the plants can main-
tain an upright orientation, the new modules appear well
below the previous tier, at the base of the plant at ground
level.
Older plants of T. anomalum may thus display two dis-
tinct kinds of branching, one consisting of 2-part tiers
around an inflorescence, and one resulting from the emer-
gence and development of a reiteration shoot into a suc-
cessional trunk module. These two branching patterns
usually look quite different, the first with evidence of the
inflorescence in the fork of the tier and no leaves sub-
tending the branches, and the second with no inflores-
cence in the fork and usually evidence of leaves
subtending each branch. As the series of reiteration
shoots matures into senescing upper branches and declin-
ing lower branches, inflorescences are sometimes aborted,
and new leaves become smaller. Older leaves fall from the
plant but senescing branches are not dehiscent, and the
aged plant finally becomes a tangle of intervening
branches, an herbaceous thicket.
The Robinson (1996) formula for this architecture ap-
proximates [O]t[O]t, but architectural patterns in
Tubocapsicum are actually somewhat intermediate be-
tween several of the growth models of Halle et al. (1978),
notably Chamberlain’s, Leewenberg’s, Prevost’s and
Nozeran’s. Early growth stages resemble those described
by Bohs (1989) for Cyphomandra Mart. ex Sendtn., but
reiterative branching is usually from near the base of the
plant in Tubocapscium rather than immediately below
branch tiers as in Cyphomandra. In neither Capsicum
nor Vassobia was an initial, orthotropic phase observed,
and plants of those genera developed directly from the
eophylls to plagiotropic branching. Subsequent growth
in these two genera was continuous rather than reitera-
tive as in Tubocapsicum. Mature growth architecture of
Tubocapsicum is thus distinctive in the dominance as-
sumed by lowest branches in a tier and the usually ac-
companying reduction or senescence of the subordinate
branches.
Vegetative Histology
Leaves. The leaf is thin and dorsiventral with a weakly
differentiated boundary between palisade and spongy
mesophyll. The midrib has a prominent, rounded adaxial
ridge and a larger protruding abaxial ridge. The cuticle
is striate. Epidermal cells are flat and thin on both
surfaces.Stomata are more common abaxially. The sto-
mata (Figures 8E, F, G) are either anisocytic or
anomocytic. Guard cell pairs are subcircular and the outer
and inner stomatal ledges, or flanges, are nearly smooth.
72 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
Adjacent epidermal cells are usually three, sometimes four.
Paradermally, adaxial epidermal anticlinal walls are undu-
late to sinuous; abaxial epidermal anticlinal walls are more
strongly sinuous and very irregular. The palisade is one-
layered, with wide cells (1.5:1 l/w ratio) comprising ca. 25%
of the mesophyll thickness. Spongy mesophyll is 3-5
layered. Spongy air space in representative cross-section
is ca. 50%. The cells are irregular with short or long lobes.
Substomatal air cavities are large and extend up to the pali-
sade layer. Midrib vasculature is a broad arc with more
or less continuous xylem and strands of adaxial and abaxial
phloem. Small vascular bundles may be bicollateral.
Smaller vascular bundles have weakly differentiated
sheath cells which contain plastids and often appear simi-
lar to neighboring mesophyll cells. Occasional druses are
present at the spongy/palisade border. Crystal sand is
present, especially in midrib ground tissue.
Figure 8. Tubocapsicum anomalum (Franch. & Sav.) Makino, pollen and leaf structure. A-C, Pollen structure; E-G, leaf epider-
mal features. A, Pollen grain, LM view. Scale bar = 7 µm; B, Pollen surface SEM view.Scale bar = 2 µm; C, Pollen grain exine
section, TEM view. Scale bar = 2 µm; D, Pollen grain polar view, SEM view. Scale bar = 10 µm; E, Leaf adaxial epidermis and
stomate (LM). Scale bar = 50 µm; F, Leaf abaxial epidermis and stomate (LM). Scale bar = 50 µm; G, Leaf abaxial surface SEM
view showing stomate. Scale bar = 10 µm.
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 73
Lamina development. This description is based on ap-
pearance of tissues in transsections. The expected mo-
saic growth patterns cannot be demonstrated with this
material. The young primordium begins the expansion
of a lamina several hundred micrometers above the point
of attachment (base) of the leaf peg. The lamina is six
layers deep at inception and the eventual thickness of all
developing laminae observed is seven layers. The outer
two layers at the margin of laminal growth appear au-
tonomous and provide for the epidermal and layers im-
mediately beneath. The third layer below the margin
produces the two central layers. A few millimeters proxi-
mal to the margin, the fourth layer down from the adaxial
surface produces two layers by division of the cell line.
This was noted in all but the youngest primordia that have
a lamina.
Stem and node. The stem has a thin epidermal layer
with small, stalked glandular trichomes. Trichomes have
uniseriate bases and rounded capitate heads of about 4
cells. Beneath the epidermis is an irregular layer of an-
gular collenchyma, 3-10 layers deep. The remainder of
the cortex and pith comprise medium-sized parenchyma
cells with no idioblastic cells. Dispersed crystal sand is
common in the ground tissue, and large druse-sized clus-
ters of crystal sand are found there occasionally. In the
young stem, several millimeters below the apex, the xy-
lem in the circular-appearing siphonostele is only differ-
entiated in the leaf traces. Phloem occurs in separate
strands, both internal and external to the xylem.
The stele appears continuous and is lobed in the vicin-
ity of leaves. The stele bulges into a leaf buttress and has
three well-matured areas of xylem: a median and two lat-
eral traces arranged as a three-trace, unilacunar (3:1)
node. An examination of the nodal cross sections from
proximal to distal shows the following pattern. A bulge
of siphonostele in the leaf buttress produces a median and
two large lateral traces that become separated from a
single stelar gap. At higher levels, the gap is closed by
newly differentiating procambium that extends in the di-
rection of the leaf base to form a crescent. The crescent
surrounds the axillary bud. At a higher level where the
leaf base is entirely separate from the stem, the axillary
bud is still included within the adaxial ridge of the leaf
base. It becomes free several hundred micrometers above
the point of separation of the leaf.
Just below the level of leaf base separation, the lateral
bundles of the petiole fork at their proximal (ventral) ends
to form two small round veins that assume a marginal
position. When the lamina begins to protrude from the
edges of the primordium, these new veins become the mar-
ginal veins of the decurrent lamina.
At the shoot apex, young leaf primordia are semicir-
cular in outline and show an early onset of abaxial defor-
mation and the protrusion of a pair of laminal primordia.
After cessation of expansion of the abaxial side of the
primordium, the adaxial side undergoes periclinal divi-
sions that form the adaxial ridge.
Floral Structure
Morphology. Flowers of Tubocapsicum are similar to
those in Capsicum, Aureliana, Withania, and Vassobia,
and as in the last three, the filaments are inserted into
stapets or stirrups (Huber, 1980) which are are fused to
the corolla tube. The stapet (filament base) is appressed
to the ovary, and in Tubocapsicum, Aureliana, Vassobia,
and Withania, a groove between each stapet allows nec-
tar to flow upwards from the nectary at the ovary base.
Flowers of Tubocapsicum are distinctive in the adnation
of the filaments to the corolla tube above the stapet to as
high as the corolla sinuses. Anthers are basifixed and
versatile. When the corolla opens all anthers are bent
inwards nearly 90° forming a spoke-like shield over the
corolla entrance. Soon, they become erect again (Figures
5, 6C, E, F, G). The ovary is superior, smooth, glabrous,
and greenish white. Nectaries are the Type I of
Bernardello (1986) in which the form is hardly visible
from outside, although they do have an ill-marked yel-
lowish color that is distinct from the rest of the ovary.
The base of the style is sunken into the top of the ovary.
Gynoecium. The ovary is two-locular with no basal
cross septum to create a false four-locular appearance.
Dorsal carpel bundles continue to the top of the style. The
style, hollow in the lowermost portion, is comprised of
solid ground tissue through most of its length. Near the
tip, the style divides into 2 very short lobes that are cov-
ered with short, thin-walled, multicellular, uniseriate
trichomes. The septum/placenta is inserted at the base of
the ovary and is free from it except at the apex. At the
base it consists of a blade-like septum that bears no ovules.
Slightly above the base, the blade-like septum expands
on its two lateral faces into a pair of hemispherical or
hemiellipsoidal ovule-bearing placentas. The internal wall
of the ovary and the septum are bounded by a differenti-
ated epidermal layer, but this is not evident on the
placenta. At the apex, the placental region ends leaving
as a continuation the blade-like septum, which at this point
divides into two branches just before uniting with the apex
of the ovary wall at the position where the style is inserted.
Androecium. An Oxalate Package (OP, D’Arcy et al.,
1996) is present in the anthers of Tubocapsicum. At the
distal ends of the anther septa, just interior to the
epidermis, a series of 6-8 cells differentiate by the time
of mitosis. The walls of these cells then disintegrate, leav-
ing a cavity partially filled with calcium oxalate. The
epidermal cells bounding these cells remain small, and
at anthesis they are much smaller than the epidermis cov-
ering the rest of the anther. Anther dehiscence takes place
by rupture of these differentiated small epidermal cells.
Pollen. The pollen of Tubcapsicum anomalum was de-
scribed by Huang (1972) and Zhang & Lu (1995) as be-
ing nearly spheroidal with a mean size of 27.2 × 28.5 µm
(Figures 8A, D); 3-colporate; colpi wide, the middle parts
nearly as thick as the ends; colpus membrane tuberculate;
os lalongate, short and narrow 8.0 × 0.8 µm in size; ex-
ine 2.0 µm thick; exine fine rugulate (Figures 8B, C, D);
columellae uneven in size, foot layer nearly as thick as
74 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
the endexine (Figure 8C). Huang reported Tubocapsicum
pollen to be among the smallest of the 21 taxa of
Solanoideae he described, but in the work by Zhang &
Lu (1995) it was nearly the same as the Physalis and Cap-
sicum taxa they described. Neither study noted charac-
ters that would separate the pollen of Tubocapsicum from
the similar genera Capsicum, Lycopersicon Mill., Physalis
L., or Solanum L.
Gynoecial Vasculature. Above the divergence of the
perianth and androecial traces within the receptacle, the
remaining vasculature does not converge into a continu-
ous cylinder but continues as two ventral traces that di-
verge at higher levels. First, below the locules, the traces
move inward, and at the base of the locules they are at
the sides of the emerging placental septum. The septum
is at first nearly plane but it soon develops thickenings
on each side in the medial region which enlarge and be-
come the ovule bearing placentas. At the level of the base
of these thickenings, the two vascular traces branch in
the plane of the septum, the smaller outer branch moving
to the periphery of the ovary where it assumes the posi-
tion of the dorsal trace, soon branching again in the ovary
wall. The stronger branches of the ventral traces move
slightly inwards where they are within and near the edge
of the placental thickened region.
Near the top of the ovary, still within the placentas,
these major ventral traces branch again, this time tangen-
tially to the septum. The branches arch across the top of
the placenta and downwards into the placenta where they
branch and anastomose further, now in a downward
orientation. Above these upper placental branches, the
major traces continue to the top of the ovary. Near the
top of the ovary, the septum and placentas are separated
into two parts which fuse with the ovary wall. The two
vascular traces continue into the style, which is sunken
in the top of the ovary and is hollow basally.
In lateral view, the two major gynoecial traces branch
weakly near the base of the ovary which provides a weak
vasculature for the ovary wall. Near the top of the
placenta, the traces branch again and the branches arch
downward enervating the marginal regions of the
placentas. Continuation of vasculature in the style is quite
weak.
The floral vasculature of Tubocapsicum may be con-
sidered against descriptions of other Solanaceae by
Bernardello (1986), Murray (1945), Huber (1980), and
Moscone (1986). In comparing Murray’s survey of the
floral vasculature of the Solanaceae, the vasculature of
Tubocapsicum resembles that of Capsicum frutescens L.
in which the vasculature continues as two traces above
the divergence of the perianth traces, rather than converg-
ing into a cylinder as in other taxa she observed. It also
resembles her Physalis description, in that the two adaxial
bundles move to the center and then issue branches in
the septum or above to supply the numerous ovules on
the massive placentas.
The gynoecium of Tubocapsicum is quite like that of
Vassobia sect. Vassobia. Comparisons of Vassobia (mostly
on V. breviflora), are based on publications of Hunziker
(1984), Moscone (1986), and our own preparation of flow-
ers of V. breviflora. In V. breviflora, the vasculature con-
verges into a ring above the corolla/androecium
divergences, and from this, two traces extend into the
ovary, which with branching, extend into the style to the
stigma. However, the septum proliferates near the base
into a false septum which partitions the basal half of the
ovary into four locules. Also, the lowermost ovules are
immersed in tissue of the ovary wall, and according to
Hunziker (1984), at maturity there is a mixture of ovary
parenchyma and ovule tissues. This last feature was in-
dicated by Hunziker as significant for separating Vassobia
from Capsicum at the generic level. In Vassobia, the style
is solid throughout rather than basally hollow as in
Tubocapsicum.
Gynoecial vasculature in Tubocapsicum displays sev-
eral differences from that seen in tribe Lycieae as outlined
by Bernardello (1986). In Lycium L., Bernardello reported
extension of the two dorsal traces through the style, but
the style was solid, not hollow as appears to be the case
in Tubocapsicum judging from serial sections. In Lycium
too, the style is not recorded as being immersed in the
ovary apex. In most species of Lycium and in Grabowskia
Schltdl. the septum and placenta are supplied by four ven-
tral traces, 2 for each carpel. In Tubocapsicum and some
species of Lycium, there is only the single ventral trace
for each carpel which branches at the base of the ovary to
yield traces in the dorsal and ventral positions. Also, the
separation of the septum into two elements occurs only at
the very apex of Tubocapsicum, whereas in Lycium it
sometimes involves the top half of the ovary. The spe-
cies of Lycium with a single ventral trace per carpel, L.
ameghinoi Speg. and L. californicum A.Gray have single
ovules and are regarded as advanced in the genus
(Bernardello, 1986).
Embryogeny. Ovules of Tubocapsicum are anatropous,
unitegmic, and tenuinucellate. The integument is four
cells thick before it has surrounded the single-walled nu-
cellus at the archesporial cell stage. At this time, the vas-
cular tissue has not differentiated into the funiculus.
Finally, a vascular bundle does differentiate within the
funiculus. While the integument is still growing, there
is no nucellar beak. By late stages, no micropyle or nu-
cellar beak is distinguishable. The nucellus has produced
a thick tissue by periclinal divisions adjacent to the
funiculus. At this stage, the integument is about seven
layers thick. There is no observable micropyle at the point
where the nucellus and integument merge at the surface.
In contrast, Vassobia has campylotropous ovules. There
is neither a nucellar beak nor periclinal buildup of tissue
in the micropylar area. The micropylar canal is clearly
distinguishable back to the tenuinucellate embryo sac.
Fruit. While sclerocytes or stone cells are common in
the Solanaceae, none of any kind were found in any of
the hundreds of fruits of Tubocapsicum anomalum
examined.In Vassobia, Hunziker (1984) noted “Bayas
donde la placenta crece centrifugamente y se confunde con
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 75
el pericarpo; en este ultimo concreciones esclerosas,” but
we found no such mixing of pericarp and placentas in the
serial sections of Vassobia breviflora that we examined.
No giant cells were seen, a feature recorded by Fridvalsky
& Nagy (1966) for Capsicum and perhaps confined to that
genus.
The juicy, ripe berry is bitter to the taste. The fruit
persists as a shiny berry for more than a month and then
darkens, wrinkles and shrivels on the yellowing, tardily
thickened calyx. Fruits placed in FAA and FPA preserva-
tives retained their bright red color for more than a year.
Both fresh, juicy fruits and dried, aged fruits are buoyant
in fresh water.
Seeds. In Tubocapsicum (Figure 9E), Aureliana, and
Withania, seeds are pale straw colored and do not darken,
even after years of storage under refrigeration or at ambi-
ent temperatures. They are suborbiculate, laterally com-
pressed (lenticular) (Figure 9A), and have a cavernulous
Figure 9. Tubocapsicum anomalum (Franch. & Sav.) Makino (A, B, E, F), Withania somnifera (L.) Dunal (C), and Aureliana
fasciculata (Vell.) Sendt. (D). A, Seed cross section through testa and endosperm (LM). Curved embryo position is seen twice.
Scale bar = 500 µm; B, Testa cross section (LM) showing blind pockets (p) and fiber layer (f). Scale bar = 50 µm; C, Testa LM
cross section showing lack of pockets. Scale bar = 100 µm; D; Testa cross section showing lack of pockets. Scale bar = 100 µm; E,
Entire seed, SEM view. Scale bar = 200 µm; F, Testa surface (SEM), close up of E. Scale bar = 50 µm.
76 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
reticulate surface architecture (Figure 9F) noticeable at 10X
magnification. The hilum is inconspicuous, linear, and lo-
cated between the radicle and cotyledon lobes. It is uni-
form in color, ca. 0.8 mm long, and is not recessed nor does
it form a notch.
The testa is formed of three physically distinguishable
layers, the inner two of which can be readily character-
ized from dried seeds. The outermost layer is formed of
collapsed distal portions of large surface cells. Such cells,
at least some members of Solanaceae, are filled with
hydrophillic colloidal material. Cell walls of this layer
are very thin and cellulosic, and are illustrated by Soueges
(1907) and Edmonds (1983). These thin-walled portions
collapse as seeds dry. The second layer, which is the first
to be encountered when examining dried or cleaned seeds,
is composed of the basal walls of the large surface cells.
This layer forms durable, prominent, lignified reticulate
ridges that extend to form a periclinal layer or floor in
each islet. They are symmetrically thickened around a
compound middle lamella, which is the point of attach-
ment of the collapsed layer. The innermost, or third layer,
is a fibrous, variably thickened or opaque tissue, ligni-
fied in these specimens. Within these testal layers a
curved embryo is seen twice in seed transection (Figure
9A) and having appressed cotyledon tips. The surround-
ing endosperm ground tissue is packed with storage gran-
ules that test negatively for the presence of starch.
The diagnostic testa features of Tubocapsicum and two
putatively related genera can be summarized as follows:
Tubocapsicum anomalum: seeds are ca. 1.5 mm long,
0.5 mm thick. Cleaned surfaces show a reticulate pat-
tern of lignified ridges, undulate at the surface, and deeply
sinuous near the base. Ridge height is ca. 90 µm, the
reticulum islets ca. 150 µm in diameter. Basal sinuosi-
ties appear perforated in surface view, but testa
transsections show “perforations” to be deep, blind pock-
ets at the base of sinuosities (Figure 9B, F). Beneath the
outer reticulate layer is a single transverse layer of
tracheids, cells square in outline, forming the inner seed
coat (Figure 9B).
Withania somnifera (L.) Dunal: seeds are 1.8-2.0 mm
long, and ca. 1 mm thick. The surface pattern is reticulate,
openings having a polygonal pattern of straight anticli-
nal walls. Ridge height is ca. 70 µm, and reticulum is-
lets ca. 90-130 µm in diameter. Ridges are rounded at
the surface and heavily thickened. The inner seed coat
layer is thin and homogeneous (Figure 9C).
Tubocapsicum seed, compared to that of Withania, is
slightly smaller, has sinuous rather than polygonal
reticulations, and the reticulation walls are much thinner.
Aureliana: (A. velutina Sendt. and A. fasciculata (Vell.)
Sendt): seeds are 2.0-2.6 mm long and 0.6-0.7 mm thick.
The outer testa is variably reticulate with ridges formed
of straight heavily thickened walls in A. velutina and nar-
row walls showing sinuosities and blind pockets in A.
fasciculata. Ridge height in A. fasciculata is ca. 110 µm,
with islets being ca. 220 µm diameter. Ridge height is
50-60 µm in A. velutina and islets to 142 µm diameter.
The inner testa wall is not lignified and possesses no vis-
ible cellular structure (Figure 9D). Tubocapsicum seed,
compared to that of Aureliana, is smaller, and has blind
pockets and sinuosities similar to those of A. fasciculata,
as well as comparble ridge height and islet size. Seed of
Aureliana velutina is quite different, with smaller, polygo-
nal islets and lower reticulate walls.
Axelius (1992) illustrated testa patterns in many spe-
cies of Physalis and Chamaesaracha (A. Gray) Benth.
The seeds of P. heterophylla Nees, P. angulata L, and C.
crenata Norhona that she illustrated are morphologically
similar, and the testa patterns are very close to those de-
scribed above for Tubocapsicum. In fact, several species
have the “very characteristic holes through the bottom
thickening” (Axelius, 1992), which we interpret as blind
pockets, based on our sectioned samples.
In the key of Gunn & Gaffney (1974), the seeds of
Tubocapsicum follow the same sequence of couplets as
Solanum torvum Sw. They differ from the species of Cap-
sicum presented in the key by their much smaller size,
and by the hilar area not being drawn out into a promi-
nent point or knob. They differ from these authors’ other
species of Solaneae (except Lycium) in the embryo being
seen only twice in cross section, not three times (imbricate
or coiled). In relation to Vassobia breviflora, the seeds
of Tubocapsicum are slightly smaller; have higher, nar-
rower reticulation walls; and much larger, more open re-
ticulation islets. The first wall layer, seen in section, is
also much larger in Tubocapsicum. Blind pockets are ab-
sent in Vassobia.
Seeds of the genera of Solanoideae compared in this
study share several common architectural features, includ-
ing overall shape, position of the embryo, layering of the
testa wall, and the surface reticulations of the second wall
layer. Some characters, however, such as the presence of
blind pockets, can be used for generic diagnosis. The com-
parisons provided above record some observable charac-
ters for use in assessing generic relationships, but they
are insufficient by themselves to place Tubocapsicum in
an evolutionary context with putative relatives. Aureliana
and Withania may well prove to be close allies of
Tubocapsicum, but the many similarities between
Tubocapsicum and other members of the subfamily as seen
by us and recorded by Axelius (1992) argue for caution
in espousing this conclusion.
Cytology
Tubocapsicum anomalum is reported (Walker, 1976)
to have a meiotic chromosome number of n = 24. The
diploid number 2n = 12 is usually recorded for other mem-
bers of subfamily Solanoideae.
Cultivation Notes
Seeds displayed no dormancy, showing a high rate of
germination after immediate planting from ripe fruits or
after cleaning and dry storage for many months. Seeds
germinated in about 24 days.
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 77
In the seasonal climate of St. Louis, Missouri (38° N
latitude), plants began to flower after about four months,
and bloomed continuously for several months. Several
specimens planted out-of-doors in June in beds with to-
matoes and Physalis alkekengi L. soon withered and
perished, displaying symptoms of heat/light stress, al-
though the other species accompanying them flourished.
At cooler times of the year, plants cultivated in near-full
sun and partial shade showed no differences in appear-
ance or performance, but significant differences were seen
when plants were held under 55% shade. These plants
appeared very healthy with darker green, somewhat
smaller, more broadly ovate leaves, and more compact
growth, but inflorescences remained arrested at a juve-
nile stage, and few flowers appeared.
Fruit set was high under insect-free conditions, the time
from anthesis to mature fruit being about six weeks. Fruits
did not abscise and fall but persisted on the plants sev-
eral months. After about four weeks they begin to shrivel
somewhat but remain bright colored, and usually are ul-
timately shed with the pedicel. Sometimes, however,
pedicel and shriveled fruit persist for many months.
From these observations, we presume that, in nature,
occurrence of the species is limited by high temperatures
and by excessive shade. The distribution map suggests
that the species grows only in areas of mild winters. Our
greenhouse experience suggests that Tubocapsicum
anomalum also requires an abundant and continuous sup-
ply of water. When plants went unwatered for 2-3 days,
they usually fell over, but they would return to an erect
posture 1-3 hours after watering.
Nectar and Pollination. No pollinator observations
are known. The size and shape of the flowers suggest
small insects, possibly bees or flies, as vectors based on
the dictum that “flowers borne singly are generally larger,
but not enormously larger, than the insects that pollinate
them.” (Proctor et al., 1996: 41). No scent was detected
from the flowers, even after they had been placed in new
glass vials for several hours to accumulate any scents
present.
Nectar is sometimes copious, and sugar hygrometer
measurements usually indicated sugar concentrations ex-
ceeding 55%. Such high concentrations are suggestive
of bee pollination (Proctor et al., 1996: 41). The specific
nectar components and their relative abundance were
found to be: fructose 9.3-10.6%, glucose 7.4-8.8% and
sucrose 80.6-83.3% (C.E. Freeman, pers. comm.). High
sucrose to hexose ratios [sucrose/(fructose+glucose)] such
as the 4:1 found in Tubocapsicum anomalum are notably
attractive to the mainly nocturnal moths, and to diurnal
butterflies and long-tongued bees (Proctor et al., 1996:
41). Flowers of Tubocapsicum anomalum remain open
day and night, which is unusual in Solanaceae.
Discussion, Relationships
Tubocapsicum is clearly a member of the Solanaceae
subfamily Solanoideae, sharing such characters as discoid
seeds, curved embryo, baccate fruit, floral actinomorphy,
chromosome number x = 12, and others noted by D’Arcy
(1975, 1991) and Hunziker (1979). These characters ex-
clude it from the other subfamily, Cestroideae. However,
the relationship of Tubocapsicum within the Solanoideae
is not clear from morphological evidence. Tribal arrange-
ment in the Solanoideae is still poorly defined, but the
works by Hunziker (1979), D’Arcy (1975, 1991), and
Benítez & D’Arcy (1997) recognize eight tribes
(Capsiceae, Datureae, Hyoscyameae, Jaboroseae,
Juanulloeae, Lycieae, Nicandreae, Physaleae) in addition
to the tribe Solanaeae which remains quite heterogenous.
Fusion of calyx lobes and accrescence of the calyx in
fruit are a common characteristic of the Solanoideae, but
the degree of fusion varies greatly and has been used in
assigning taxonomic relationships. In tribe Capsiceae,
including Aureliana, Lycianthes (Dunal) Hassl., Vassobia,
and Witheringia, flowering calyx lobes are fused for much
of their length (complete prefloration), and floral egress
is by stretching (D’Arcy, 1986; Benítez and D’Arcy,
1997). However, in Tubocapsicum anomalum (Figure
3D), the mature calyces remain small at the base of the
fruit in most specimens.
In tribe Physaleae, which embraces Physalis, Withania
and other genera, the calyx is lobed, and floral egress re-
quires little or no stretching. In fruit, the calyx is
accrescent, enveloping the fruit. Corollas in both the
capsicoid and physaloid genera generally have a short tube
that is basally appressed to the ovary, a more or less cam-
panulate limb, and short lobes, although in some, the
lobing is more than halfway down the limb. The calyx
of Tubocapsicum anomalum is of the capsicoid form: the
lobes are fused around the bud leaving only a minute api-
cal opening for floral egress, and the body of the calyx is
not accrescent after anthesis. Thus, the calyx and other
features of the plants are highly suggestive of a relation-
ship with Capsicum and other similar genera.
However, in Tubocapsicum, where corolla and adnate
filaments are appressed to the ovary, there are grooves
between the filaments that permit nectar release. Such
grooves are also found in Aureliana, Withania and per-
haps other genera, but not in the species of Capsicum that
we examined. In Tubocapsicum and Aureliana, the stapet
(or stirrup), where the filaments are inserted, is contin-
ued into small teeth. This condition was not found in the
other capsicoid species or Withania species we examined.
Tubocapsicum differs conspicuously from other mem-
bers of the Solanoideae in the adnation of its filaments to
the corolla limb. Diagnostic for Tubocapsicum is the con-
tinuation of adnation of the filament above the level (or
what corresponds to the level) of filament insertion in
other groups, so that the filament is adnate to the corolla
limb, from the base of the tube to the base of the sinuses
in the limb. Such adnation of the filament up into the
corolla limb was not seen in other putatively related plants.
The recurved corolla lobes of Tubocapsicum are also
distinctive. Tubocapsicum differs conspicuously from
Capsicum in lacking giant cells of the ovary as reported
by Fridvalsky & Nagy (1966).
78 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
In their analysis of chloroplast DNA, Olmstead et al.
(1999) found Tubocapsicum, Withania, and Aureliana as-
sociated on a clade well separated from others in their
inferred phylogeny of the family. Relationships among
the three genera are not particularly close (weakly
supported, Olmstead, pers. comm.), and their geography
is disparate. Withania ranges from the Canary Islands to
the Himalayas, Aureliana is restricted to Argentina,
Bolivia, and Brazil, and Tubocapsicum is restricted to east-
ern Asia. This study of Tubocapsicum is not concordant
with the DNA-based conclusions, but examination of a
larger sample of species from similar genera in the
Solanoideae may further illuminate the phylogeny.
Taxonomic Treatment
The first species of Tubocapsicum was described in
1879 by the French botanists Franchet and Savatier, as
Capsicum anomalum. Over the years, it was described
again several times under different names, as noted below.
When von Wettstein (1898) treated the Solanaceae for
Engler & Prantl’s treatment of the world’s plants, he sepa-
rated C. anomalum from other species of Capsicum as
“Sect. II. Tubocapsicum” with only the comment, “flower
broadly campanulate.” Then in 1908, the Japanese bota-
nist Makino, elevated Wettstein’s name to the rank of
genus, and he transferred the single the species, Capsi-
cum anomalum, into Tubocapsicum.
Georg Bitter (1913, 1917) placed two names, Solanum
anodontum and Solanum philippinense, into synonymy
under Capsicum anomalum, and outlined the geographic
range of the genus. Bitter did not concur with Makino’s
ranking of Tubocapsicum as a distinct genus and followed
Wettstein in ranking it as a section of Capsicum. Since
Makino’s establishment of the genus and Bitter’s com-
ment on it, there has been scant reference to
Tubocapsicum. Hunziker (1958), who had not seen ma-
terial at the time, considered it to be a section of Capsicum.
Numerous non-descriptive references to the plant are
found in Asiatic floras, and short descriptions have ap-
peared in a few floras: Makino (1963) and Ohwi &
Kitagawa (1983) for Japan, Liu & Ou (1978) and D'Arcy
& Peng (1998) for Taiwan, and Kuang & Lu (1978) for
mainland China.
Additional limited information on Tubocapsicum has
been published by: Zhang & Lu (1995); Franchet &
Savatier (1875): Makino (1926: 7; 453); Liu & Ou (1978:
548); Kitamura (1937: 18); Walker (1976: 919); Huang (1972:
227) (pollen); Makino (1963: 545) (in Japanese); and Ohwi
& Kitagawa (1983: 1317) (in Japanese). The only illustra-
tions of the plants that we know of are: Savatier (1874);
Liu & Ou (1978: 549); Wu & Raven (1998) and D'Arcy &
Peng (1998). The text and illustration from Zhang et al.
(1994) can be seen on the World Wide Web at http://
herbaria.harvard.edu/china/solan/tubocaps.htm. Pollen
was treated by Zhang & Lu (1995) and leaves and seeds
by Zhang & Wen (1996), and these authors also illustrated
pollen, leaf surfaces and seeds.
All except Liu & Ou (1978) are in the language of the
country treated, and in these works, Tubocapsicum has
usually been recognized as a genus. Walker’s English-
language Flora of Okinawa (1976), an exception, recog-
nized the species under Capsicum.
TUBOCAPSICUM (Wettst.) Makino, Bot. Mag. Tokyo 22:
18.1908.
Capsicum sect. Tubocapsicum Wettst. in Engl. & Prantl,
Nat. Pflanzenfam. IV(3b): 21. 1891.—TYPE: T.
anomalum (Franch & Sav.) Makino.
Description
Herbs, glabrate, the trichomes simple. Leaves
alternate, simple, subentire, ovate, secondary veins
arcuate, brochidodromous. Inflorescences few-flowered
fascicles in the upper leaf axils, sometimes subtended by
a small, early-deciduous, leaflike bract; peduncle wanting;
pedicels only slightly expanded upwards, glabrous. Flow-
ers 5-merous; prefloration almost complete (the budding
perianth fused almost to the top, enclosing the inner parts),
in bud the calyx showing 5 minute terminal lobules and
a minute hole or port; calyx cupular, apically truncate,
with only the slightest evidence of five terminal points,
hardly accrescent; corolla oriented with the mouth facing
downward, glabrous outside, actinomorphic, the tube
short, included in the calyx, the limb broadly campanulate,
glabrous, lobed 1/4 its distance, the lobes deltoid or obtuse,
recurved, with no plicate area evident after opening; sta-
mens inserted at the base of the corolla limb; the filaments
basally broad, thinner upwards, narrowed into a slender
neck just below the anther, each with 2 teeth or umbos
just above the point of insertion, adnate to the corolla tube
a short distance above the insertion, free from one an-
other with narrow grooves between them, the filaments
at first bent outwards at the top of the corolla tube and
adnate to base of limb before arching upwards to the co-
rolla mouth; anthers half-exserted from the corolla, del-
toid-oblong, ca. 1 mm long, dehiscing laterally but the
pollen sacs (thecae) opening inwards (ventrally,
adaxially), the ventral sacks folding flat against one
another; ovary included in the filament expansions, the
lower half differentiated into a copious-flowing yellow
nectary to which the filament expansions are applied, the
style straight, terete. Fruit a subglobose or ellipsoidal
berry (Figure 4D); seeds pale yellow, discoid, favose.
(Figure 9A, B, E, F).
Tubocapsicum includes the widespread species T.
anomalum, and populations from near the sea on the
southeastern end of Shikoku Island, Japan. The Japanese
specimens were recognized by Makino as T. anomalum
var. obtusum and as the species, T. obtusum, by Kitamura.
Key to the Species of Tubocapsicum
1a. Leaf apices acute or acuminate; fruiting calyx apically
contracted, the thin, distal sleeve region not
accrescent, inconspicuous, mostly free from the berry;
seeds mostly 1.4-1.7 mm across ......... 1. T. anomalum
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 79
1b. Leaf apices obtuse and short acuminate; fruiting ca-
lyx not contracted apically, the thin apical sleeve area
slightly accrescent and applied to the base of the berry;
seeds mostly 1.5-2 mm across .............2. T. obtusum
1. Tubocapsicum anomalum (Franch. & Sav.) Makino, Bot.
Mag. Tokyo 22: 19. 1908. Capsicum anomalum Franch.
& Sav., Enum. Plant. Jap. 2: 452. 1879. SYNTYPES:
Japan, Tanaka (P), Hakone and Nikko, Savatier 2166
(P). A suite of five specimens, seen and cited below,
was annotated as collected by Savatier, 2 from
Hakone, 3 from Nikko, but the number 2166 does not
appear on any of them. A specimen labeled Tanaka,
Savatier 2166 (K ex L), might be taken as the holotype,
but it is labelled as “Capsicum cordiforme Mill. var.
truncata Miq.,” and we strongly doubt that it was ac-
tually seen or collected by Savatier. LECTOTYPE,
designated here: in tractu Nikko, Savatier s.n. (P). The
specimen chosen as the lectotype has a packet or small
envelope mounted on it, upon which there is a sketch
of a stamen inserted on the corolla and “Capsicum!
(anomal.).” We believe that this specimen is of the same
gathering as that cited by Franchet and Savatier in the
protologue, and that the writing on the packet may be
the work of Georg Bitter. The specimen bears a stamp
saying “Donné par M. Franchet, juillet 1882,” and it
was annotated by Bitter.
Solanum anodontum Lévl. & Vaniot, Le Mond Pl. 10: 37.
1908.—TYPE: Korea, Cheju Do (Quelpaert), Urb.
Faurie 776 (E).
Solanum philippinense Merr., Philip. J. Sci. 7: 351.
1912.—TYPE: Philippines, Luzon, Williams 1275 (K).
Capsicum boninense Koidz., Fl. Symb. Orientali-
Asiaticae. 3: 1. 1930. Tubocapsicum benignancy
(Koidzumi) H. Hara, Enum. Sperm. Japan. 245.
1948.—TYPE: Bonin Is., Wright 200 (K, P, US).
?Capsicum minimum auct. non Roxb: Henry, Trans. Asiat.
Soc. Japan 24 Suppl. 65. 1901. Henry’s report is based
on Henry 356 (MO).
Solanum biflorum auct. non Loureiro: Savatier in Iinuma,
Somoku-Dzusetsu, ed. 2, 3: 47. 1874.
Capsicum cordiforme ?Mill. var. truncata Miq. in
sched.— TYPE: Japan (K, P). Bitter (1917) referred
to a specimen in (BREM), probably a duplicate of ones
we cite below, annotated as Capsicum cordiforme Mill.
var. truncata Miq. We have not been able to verify
the valid publication of this varietal name.
Description
Erect herbs to 1.5 m tall; first stems stout, soon terete,
later stems weak, green, terete, glabrate, drying ridged,
the few trichomes short, simple. Branching mostly in di-
chotomous tiers. Leaves alternate, simple, subentire,
ovate, 7-40 cm long, apically acuminate, basally obtuse
or acuminate and narrowed into the petiole, sometimes
somewhat oblique, the margins sometimes somewhat
wavy, membranous, glabrate, with sparse minute simple tri-
chomes scattered on the lamina and vein, the midvein be-
neath continuous with the petiole, lateral veins arcuate,
mostly 5-6 on each side, brochidromous. Inflorescences
few (to 6)-flowered fascicles in the upper leaf axils, some-
times subtended by a small, early-deciduous, leaflike bract;
peduncle wanting; pedicels ca. 5 mm long, only slightly
expanded upwards, glabrous. Flowers 5-merous;
prefloration almost complete, in bud the calyx showing 5
minute terminal lobules and a minute hole or port; calyx
2.5-3 mm long, glabrous, cupular, apically truncate, with
only the slightest evidence of five terminal points, not
splitting, not accrescent; corolla oriented with the mouth
facing downward, opening uniformly yellow, later becom-
ing brownish, almost waxy or shiny, glabrous outside,
actinomorphic, subglobose or turbinate in bud, the tube
short, not exserted from the calyx, the limb broadly
campanulate, glabrous, 3-6 mm long, the mouth ca. 6 mm
across, lobed 1/4 way down, the lobes deltoid or obtuse,
recurved, minutely puberulent adaxially, the margins
granular with minute trichomes on the edges, and no pli-
cate area evident after opening; stamens inserted at the
base of the corolla limb, glabrous, the filaments 2.5 mm
long, basally broad, thinner upwards, each with 2 teeth
or umbos just above the point of insertion, adnate to the
corolla tube a short distance above the insertion, free from
one another with narrow grooves between them, the fila-
ments at first bent outwards at the top of the corolla tube
and adnate to the base of the limb before arching upwards
to the corolla mouth, narrowed into a slender neck just
below the anther; anthers half exserted from the corolla,
deltoid-oblong, pale yellow, 1 mm long, not apiculate, de-
hiscing laterally but the pollen sacs (thecae) opening in-
wards (ventrally, adaxially), the ventral sacks folding flat
against one another, the dorsal sacks held apart by the
broad shiny anther dorsum; ovary glabrous, included in
the filament expansions, the lower half differentiated into
a copious-flowing yellow nectary to which the filament
expansions are applied, the style white, straight, terete,
uniform in diameter, 2.5 mm long, inserted in a short de-
pression in the ovary apex, the stigma minute, green,
capitate, situated at about the level of the anthers. Fruit
a subglobose or ellipsoidal shiny scarlet juicy berry, long
persistent on the plant, 9 × 7 mm, the pericarp thin,
translucent, the endocarp scarlet, soft and juicy, partly im-
mersing the abhilar edges of the seeds; calyx not
accrescent; fruiting pedicels somewhat elongating but not
thickening until extreme age; seeds, pale yellow, discoid,
favose, 1.7 × 1.4 × 0.6 mm (Figure 9).
Distribution and Phenology
Tubocapsicum anomalum ranges (Figure 1) from the
Japanese island of Honshu, south through the Bonin and
Ryukyu Islands, Taiwan, the Philippine Islands and as far
south as eastern Borneo. From this northeast-southwest
axis (from 38°N to the equator), it ranges westward to
the Korean island of Cheju Do, and into China between
the Huang He (Yellow River) and Hongshui River.
The species occurs from sea level to 2,100 m elevation
(Mt. Mukwashan Forest, Shoufeng District, Taiwan). North
80 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
of 27°N latitude, all collections seen were below 900 m.
Merrill (1923) recorded its occurrence in the Philippines in
“mossy forest, 1,800 to 2,200 m, and along streams in
shaded ravines at low and medium altitudes.”
In Asia, most flowering specimens have been collected
from August to November, but some were made as early
as March. Most fruiting specimens were made between
September and November, but at least some have been
made in nearly every month.
Herbarium material of Tubocapsicum anomalum is
commonly misidentified as Capsicum annuum, Lycianthes
spp., or Solanum spp. Capsicum annuum can be distin-
guished by elongate and mostly bluish anthers; in
Lycianthes and Solanum, the anthers have terminal pores.
Specimens of Tubocapsicum with fruits but lacking flow-
ers in particular to have been confused with these genera.
Capsicum annuum can be distinguished by its seed that
are mostly more than 3 mm across in contrast to the 2
mm or less of Tubocapsicum anomalum seeds, and the
fruiting calyx that is usually much larger. Pedicels of most
Capsicum specimens from Asia are straight and stiff,
while those of Tubocapsicum are curved and flexuous.
The fruiting calyx of Tubocapsicum lacks the evident
nervature of Lycianthes calyces, and the evident lobes of
the Solanum species that occur in Asia.
Common Names: Chinese “Long zhu (Dragon Ball)”
(Zhang et al., 1994). Japanese “Hadaka-h_dzuki (Naked
Balloon)” (Savatier, 1874; Anon.a, 1887: 158; Anon.b, 1895:
126; Yokohama Nursery, 1908; Kawakami, 1910); Tachi-
hiyodori (Standing brown-eared bulbul.), Yama-h_dsuki
(Mountain or Wild Balloon) (Anon.b, 1895: 126).
Specimens Examined. BORNEO. N. Borneo, Lahad
Datu Dist., Kennedy Bay, D. Brand SAN 20102 (LL). SW
Borneo, Kuntim Lian?, Batu Leabi, Winkler 2734 (LL).
CHINA. FUJIAN: Jianning Xian, Li 10551 (PE);
[Fukien] Siong-gu-liang, Hok-Chang, Tang Siu Ging
16367 (A, MO); Wuyi Shan, 1,100 m, Wuyi Shan Team
80-0634 (MO). GUANGDONG: Lechang Xian, Chen
42079 (PE); Liannan Co., Li 414 (MO); Renhua Xian,
900 m, Liang Deng 7916 (PE); Shixing Co.,
Zhangdongshui, 400 m, Xing & Ye 942 (MO); Xinfeng
Xian, 700 m, Liang Deng 7916 (PE); Yangshan Xian, 420
m, Guangdong & Guangxi Exped. 0383 (PE); Zijin Xian,
450 m, Wei 120883 (PE).GUANGXI: Damiaoshan Co.
Luo Tong Xiang 15680 (MO), Sangfang Qu, Pingshi
Xiang, Luo Tong Xian, Chen 15680, 15735 (both MO);
Antai Qu, Xiaoshang Xian, 600 m, Chen 17319 (MO);
Dayao Mt., Lu 4492 (PE); Without other locality,
Longshen Exped. 50396 (PE); Witsap District, Tou Ngok
Shan near Tung Chung village, Tsang 23227 (P);
Yongshui Xian, Chen 15680 (PE); Damiaoshan Co.,
Sanfang Qu, Pingshi Xiang, Chen 16590 (MO); Xingan
Xian, 800 m, Yu 9000242 (PE); (Other counties (“Xians”):
Jinxiu, Rongschui Xinan, fide Zhang Zi-yun). GUIZHOU:
Fanjing Mountain, Sino-British Plant Exped. 38 (K),
Fanjingshan Mt., Guiyang Teacher College F0038 (PE);
Bijie Xian, 1,430 m, Yu 857 (PE); Jiangkuo Xian 800 m,
Bartholemew et al. 146 (PE); 950 m, Bartholemew et al.
537 (PE); Jianning Xian, Li 10551 (PE); Leigong Mt.,
Qiannan Exped. 3800 (PE); Songtao Xian, 600 m,
Bartholemew et al. 2342 (PE); Yongjing Xian, Qiannan
Exped. 3379 (PE); (Other counties: Zunyi, fide Zhang Zi-
yun). HAINAN: without other locality, Wang 35210 (E);
HUNAN: Chili Xian, 600 m, Hunan Exped. 636, 647
(both PE); Mang Shan, Pinkeng, 200 m, Huang Maoxian
112942 (MO); Xiangyi Xian 700 m, Liu 10294 (PE);
Xinning, Ziyunshan, 1,150 m, Liu 15116 (PE); 1,100 m,
Zhen-yu et al.1791 (MO). JIANGXI: Anfu Xian, 570 m,
Yue 3098 (PE); Daganshan, Fenyi City, Yao 9262 (K,
MO), 9264 (MO); Dexing Xian, without collector 5233
(PE); Guangchang Xian, Hu 5249 (PE); Jiangan Xian,
300 m, Shu-kun Lai 784 (PE); 400 m, Lai 1518 (PE);
Jinggang Mt. 1,000 m, Jie Xiong 2763 (PE); Jiulian Shan,
510 m, Xie 93167 (MO); Lichuan Co., (Flora of
Chien-Xi), near Lan-Uk village, Nungan District, Lau
4701 (US); Wuning Co., Wuning Yan, 300 m, Tan 941123
(MO); Xunwu Xian, Yue 2240 (PE); (Other counties:
Jing’an, fide Zhang Zi-yun). YUNNAN: Zhenxiong Co.,
fide Wu (1984: 1565). XIZANG (Tibet): Between Sikkim
and Bhutan, Chumby-Valley, Searight 261 (VRSL, not
located, B, destroyed, cited by Bitter, 1917). ZHEJIANG:
Changuha Xian, He 26350, 29803 (both PE), without col-
lector 30428, 30623 (both PE); Chunan Co., Jinzhijian,
Maanli, Chiu 2138 (MO); Jian Co., Longwang Shan,
Mafengan, Chiu 3305 (MO); Jin Xian, without collector
9221 (PE); Kaihua Xian, without collector 29924 (PE);
Souchang Co., Xihuasha Zhou, Chiu 671 (MO), without
collector 29568 (PE); Tianmu Mt., 550 m, Zheijiang
Exped. 29022 (PE), without collector 25353, 31270 (both
PE); Taishun Co., Wuyanlin, Chang 8661 (MO); Xianju,
Shisandu, Zhongkeng, Zhejiang Bot. Resource Team
29966 (MO); Zenhai Co., Ruiyan Shi, Chiu 133 (MO),
without collector 2043 (PE); (Other counties: Jiande,
Lin’an, Yinxian, fide Zhang Zi-yun). TAIWAN. Sakahen,
Gressit 460 (K, LL); Mt. Taiheizan, Suzuki 275 (MO);
Namba et al. C 557 (TI). CHIAYI HSIEN: Shihcho, 400 m,
Yen 6073 (HAST); Tapu Hsiang, Tingpinglin, 600-700 m,
Yen 6604 (HAST); Chuchi Hsiang, Shihcho, Yen 6873
(HAST, MO). HSINCHU HSIEN: Wufeng District, en
route from Tuchang to Kuanwu ca. 16.5 km S of Tuchang,
1,520 m, Peng 9364 (HAST). HUALIEN HSIEN:
Shoufeng District, Mt. Mukwashan Forest, Halun Station.
2,000-2,100 m, Peng 7270 (HAST, MO). Hsiulin Hsiang,
from Kuailin to Chilai, 1,425-2,160 m, Ho et al. 679
(HAST). Shuiyuan to Luanschan NW of Hualien, 1,600-
1,900 m, Tamura et al. 21517 (E). Ta-Yu-Ling, Chuang
& Kao 4373 (LL). ILAN HSIEN: From Chi-lan Forest
Station to Yuen-yang Lake Natural Reserve, 500-1,500 m,
Boufford et al. 19413 (MO). Fu Shan Botanical Garden,
700-720 m, Peng 3878 (HAST). Fu Shan, 650 m, Bufford
et al. 25099 (MO). Between Kungliao and Hutzupei (a
stone tablet), en route from Kungliao to Tali, 40 m, Peng
13566 (HAST). Nanhutaschan, Nanshan-Kiretei, 1,600 m,
Yamazaki et al. 737 (MAK-142083, TI). MIAOLI HSIEN:
Nanchuang Hsiang, along bank of Fengmeihsi, 1,500-1,700
m, Liu 339 (HAST, MO). NANTOU HSIEN: Hsinyi Hsiang,
Petyushan, cut from Shalihsienhsi Forest Road through a
D’Arcy et al. — The genus Tubocapsicum (Solanaceae) 81
valley up to a mountain ridge 1,300-1,700 m, Chang 86
(HAST). Between Chi-tow & Sunkinksea, 1,100-1,600 m,
1983, Peng 5396 (HAST, MO). Hsinyi District: Shenmu,
Peng 10975 (HAST). Luku Hsiang, Hsitou. Peng 6983
(HAST), Lin 623 (HAST). Chitou to Fenghuangshan,
Chen 231, 779 (both HAST). En route from Tungpu Hot
Spring to Rainbow Waterfall. 1,200-1,400 m, Peng 8228
(HAST, MO). Feng-shan, Chi-tou, 1,400 m, Yamazaki 160
(TI). PINGTUNG HSIEN: Bankinsing, Henry 356 (MO).
Tahanshan, Amenotaxus formosana Preserved Area, 1,300-
1,500 m, Chaw 806 (HAST). Chunjih Hsiang, along Tahan
Forest Trail from Shitiliao to Tahanshan, 500-1,300 m, Yen
et al. 572 (HAST); Wutai Hsiang, en route from Wutai to
Payutzu (Little Ghost Lake), entrance of Ching-Chieh-Shan,
Lin 306 (HAST). TAICHUNG HSIEN: Sasaki 6 (TI). Hop-
ing Hsiang, Wushihkeng, from The Low Altitude Experi-
mental Station, Taiwan Endemic Species Research Institute
to Wushihkenghsi, 1,000-1,250 m, Shen 218 (HAST).
TAIPEI HSIEN: Wulai Hsiang, Fushan Village, Kuaishan,
1,350-1,590 m, Liao 1005 (HAST); Wulai, 150-200 m, Peng
10008 (HAST, MO). Pinglin Hsiang, Taiyuku, from
Chinkualiaohsi (stream) to Preserve of Keteleeria
davidiana, 210-350 m, Liu 4 (HAST, MO); Mt.
Chihsingshan, en route to Mr. Yu You-jen Graveyard, 800
m, Peng 9879 (HAST, MO); Yangmingshan National Park,
Tatunshan mountain from the parking lot of Tarun Nature
Park to Erhtzuping, 825-840 m, Shen 273 (HAST).
TAITUNG HSIEN: Orchid Island (Lanyu) trail up to Tein
Tsiu, 325 m, Lammers 8543 (MO); Green Island (Lutao).
Forested drainage ditch leading to the sea by Hot Spring,
Peng 7616 (HAST); Along Yeh-yin stream from Yung-
hsing-nung-chuang to Ao-pen-ling, 100-250 m, Leu 1378
(HAST, MO). TAOYUAN HSIEN: Lalashan Nature
Preserve, NE of Shang Paling, along trail, entrance to
Lalashan Mt. peak, 1,550-1,700 m, Peng 14619 (HAST).
INDIA. ASSAM: Masters (B, ex CAL, destroyed, not seen,
cited by Bitter 1917). JAPAN. Without locality or collector.
Capsicum cordiforme Mill var. truncata Miq.” ex herb.
Lugduno-Batavo (K, LL, P); [-Labelled in pencil as Tanaka
Savatier 2166.] Without other locality, Bissett yr 1877 (K);
Dickins 10/81 (K); Mohnike s.n. (LL); Oldham 1861 (LL);
Tanaka 263 (P). BONIN ISLANDS: Wright 200 (K, P, US).
RYUKYU ISLANDS: Ins. Iriomote, Komi-Komidake, 50 m,
Yamazaki 17 Jun 1971 (TI); Ins. Amami-oshima, Hatusima
19988 (LL); Hatusima & Sako 23893 (LL); Ins. Eshaki
Shima, Fosberg 37371 (LL, US); Ins. Ishigaki, Mt. Banna,
Iwatsuki et al. 893 (LL, MO); Ins. Okinawa, without other
locality, (MAK-97977). Ryukyu University Experimental
forest, Yona, Kunigami-son, Tamaki 12 Oct 1957 (MAK-
141633). Amano 6858 (US). Moromizato, 31 Oct 1957 (US).
Tanyoeake, Walker 83183 (LL, US). Common but scat-
tered at seaside, Tawada 2226 (US). Elliot 1081 (US); Ins.
Yugashima, 170 m, Kanei et al. 10084 (TI); Ins. Yonakuni,
Furuse 2501, 4179, 4196, 4524, 4525, 4643 (all K).
KYUSHU. Bungo, Takenouchi 2776 (US); Segiri, 100 m,
Yamazaki 7001 (TI); Yakushima, Nagata, Kuniwari-dake,
400 m, Yamazaki 6923 (F, TI); -note rounded leaves. Yaku-
cho along river Nakamagawa, Chizu Chuma, 4 Mar 1968
(TI); Pref. Oita, Mt. Kujusan (MAK-97972). Kujusan,
Jusan-magarari, Yobikogawa, Togashi 7356 (TI);
Kagoshima: Shiroyama, Kagoshima City (MAK-97954);
Goino, Hirakawa, Murata 16044 (TI); Kumamoto:
Fukabayama, Aso-machi, Aso-gun (MAK-75361). Aso,
Takamoricho, Takamori-toge, 800 m, Yamazaki 7 Oct 1988.
(TI); Mii Sagamigawa, Oct 1886 Bisset 3822 (E). HONSHU.
Without other locality, ?Yokacku, Bisset 928, 2224 (both
E); Nikko, 1873, Savatier (P-2); Hiroshima, Miyajima in Aki,
Togasi 31 Oct 1955, TNS 1259 (E, K, TI, US); Mt. Amagi,
Suzuki 18 Jul 1980 (US). Prof. Mt. Rokkosan, Kobe City
(MAK-97966). Yamakita, Yamakita-cho, Ashigarakami-gun,
(MAK-75359); Imaizumi, Kamakura City, Kobayashi 3176
(3 Nov. 1983) (MAK-212869); Enoshima, Fujisawa City
(MAK-75357); Hakone, Hakone-machi, Ashigarashimo-
gun.Makino, Oct 1924 (MAK-97965); Hakone, Savatier (P-
3); Mt. Takatoriyama, toward the east, Zushi City,
Momiyama s.n. (MAK-234896); Mt. Takatoriyama,
Yakushido, Yusaka, Zushi City, Momiyama s.n. (MAK-
234756); Yokohama, Maximowicz. Iter secundum (LL, P);
Amano 8 (MO). Kifune, N of Kyoto, Murata 18647 (E,
LL, P, US); Satomi 9414 (MAK-236457); Takahashi 431,
(MO); Tokyo, Sep 1910 without collector (E); without
collector, (E); Azusawa, Itabashi-ku, (MAK-97955);
Inokashira, Mitaka City, Makino, s.n. (CAL, MAK-75354);
Makino 75355 (LL); Prov. Musashi: Tabata, Aug 1910
without collector (E, US); Prov. Musashi: Tabata, Aug.
1910 without collector (E); Tokyo, Tahata-Mura, Tabata,
Aug 1910 without collector (US); Musashi (Oji), 21 Aug
1908, s. coll. (VRSL); Yokahama Nursery, July 1808 (E); Pref.
Yamaguchi, Mt.Tokusagamine, Abu-gun, (MAK-130188).
Pref. Nara: Takahashi 774, (MO); Pref. Shizuoka, Amagi
Pass, Mt. Amagi, 800 m, H. Kaman 18 Sep 1961 (TI).
SHIKOKU: Ehime: Izushidera, Satomi 9088 (MAK-
236454); Mt. Iwayayama, Mikawa-mura, Kamiukena-gun,
(MAK-97970); Kochi: Mt. Dogamori, Nakamura City
(MAK-97963); Iyoki: Otoyo-mura, Nagaoka-gun (MAK-
97967); Shimonanakawa, Agawa-mura, Agawa-gun, 300 m,
Murata 18864 (TI); Muroto-city, Moroto-misaki, near
seaside, Takahashi 1223 (TI); Zyadani, Kitagawa-mura,
Aki-gun, 400 m, Murata et al. 63 (TI). KOREA. Cheju Do
(Quelpaert Island) Hongno? 33°24’N, 126°24’E, Chung Ing-
Cho 3756 (F); Faurie 776 (E); Nakai 6463 (TI); Nakai 6
Jun 1913 (TI); Taquet 1145 (E); 3078 (E, K), (both B-cited
by Bitter 1917, destroyed) (Taquet 1144 (K) is a species
of Solanum). MALAYSIA. locality illegible, Winkler 2734
(K); Sabah, Ulu Segama, Argent et al. 108201 (K).
PHILIPPINES. Loher 6643 (B, ex K, B-cited by Bitter 1917,
destroyed); Bandschan, Loher 4377 (K); Mindoro,
Escritor, Jul 1913 (Bur. Sci. 21306) (K); Laguna, Mt.
Meguiling, Elmer 17592 (LL); Ins.Luzon, Elmer 6561 (K-
2, US, B-cited by Bitter 1917, destroyed); Elmer 16996 (K);
Serviñas s.n. (Bur. Sci. 16862) (MO); Benguet, Merrill
8003 (K, LL, US, B-cited by Bitter 1917, destroyed);
Mariveles Island, Manila Harbor, Day 185 (MO); Luzon,
Mt. Magras, Edaño 6279 (LL); Benquet Prov., Mt.
Tonglon, Ramos 5406 (LL); Sorsogon, Ramos Jul-Aug
1925 (Bur. Sci. 23438) (K); Ins. Mindanao, Merrill 8164
(K, LL, US, B-cited by Bitter 1917, destroyed).
82 Botanical Bulletin of Academia Sinica, Vol. 42, 2001
2. Tubocapsicum obtusum (Makino) Kitamura, Acta
Phytotax. Geobot. 6: 18. 1937. Tubocapsicum anomalum
var. obtusum Makino, J. Jap. Bot. 3(2): 7. 1926.—TYPE:
Japan, Prov. Tosa, Muroto-zaki (Torama Yoshinaga), 3
Jan 1921 (MAK-97979).
Description
Glabrous herbs, stems drying grooved, fine lenticellate.
Leaves succulent, broadly ovate, mostly 3-8 cm long,
apically obtuse and short acuminate, basally truncate or
cordate, sometimes acuminate and narrowed into the
petiole, lateral veins arcuate, mostly 3 on each side. In-
florescences few-flowered fascicles in the axils of leaf
pairs, mostly with 1-2 small (-1 cm), apically rounded
leaves in addition to the paired normal leaves; pedicels
1-2 cm long. Flowers with calyx 4 mm long, cupular,
apically truncate, the apical sleeve region 2 mm long; co-
rolla deeply lobed, the lobes narrow, acute, ca 5 mm long;
anthers oblong, 1.5 mm long, not apiculate, the thecal re-
gion light colored, the narrow connective dark. Fruit a
subglobose or ellipsoidal red or orange berry, 8-10 mm
across, the pericarp thin, translucent; calyx slightly
accrescent; pedicels somewhat elongating and thickening;
seeds, pale yellow, discoid, favose, 1.7-2.0 mm across.
The species differs from Tubocapsicum anomalum in
its smaller, thicker and more blunt-tipped leaves, in its
larger fruiting calyces, and perhaps in having slightly
larger seeds. In some collections, the fruiting pedicel ex-
pands somewhat apically, thus resembling some speci-
mens of feral Capsicum annuum, but the seeds are much
smaller than in that species. Although several specimens
were available for study, only one (MAK-44282) bore a
flower that could be dissected. The anthers are different
from those of T. anomalum, and it is probable that this
taxon is member of a different genus. The stamens and
corolla shown by Liu & Ou (1978) in their illustration of
Tubocapsicum anomalum are more like those of T.
obtusum than those of T. anomalum, in that the corolla
lobes are not recurved and the anthers lack the broad dor-
sum characteristic of T. anomalum.
Because some (or most) collections were taken near the
sea, it is possible that their succulent appearance shows
the salt-succulence commonly found in seaside vegetation
(Boyce, 1954).
The specimen we refer to as the type specimen has no
collector indication, but Makino recorded the collector as
Torama Yoshinaga, and noted that this taxon grows “at
the littoral place.” Kitamura, who transferred Makino’s
varietal name to the rank of species, indicated his doubt
that the taxon belongs to Tubocapsicum. Until additional
material becomes available, it seems best to accept
Kitamura’s disposition of these plants.
Distribution. Tubocapsicum obtusum is reported from
Japan.
Specimens Examined. JAPAN. RYUKYU ISLANDS:
Ins. Iriomote, Komi-Komidake, 50 m, Yamazaki 17 June
1971 (TI); Ins. Eshaki Shima, Fosberg 37371 (US); Ins.
Ishigaki, Mt. Banna, Iwatsuki et al. 893 (MO); Ins.
Okinawa, Yona, Kunigami-son, Tamaki 13 Jan 1970 (MAK-
128613) [depauperate]. Common but scattered at seaside,
Tawada 2226 (US); Ins. Tokunoshima, Hetono, Amagi-cho,
Sugawara 2067 (MAK-206083). HONSHU: Island of
Miakao, Koidzumi 11 Jun 1923 (US); Honshu. Pref. Chiba:
Matsudo, Matsudo City (MAK-75352). KYUSHU: Pref.
Kagoshima: Hashima, Kushikiko City, Hatusima & Sako
27989 (MAK-51647). SHIKOKU: Kochi: Shimokanoe,
Tosashimizu City, (MAK-97962); [intermediate] Muroto-
saki, Muroto City, [Torama Yoshinaga] 3 Jan 1921 (MAK-
97979, type collection).
Acknowledgements. Thanks are offered to curators of insti-
tutions who arranged for our loans of herbarium specimens (A,
CAL, E, K, MAK, P, TI, US, VRSL) as well as curators and
staff of our own institutions, HAST, MO and PE. We thank
also the Conservatoire et Jardin Botanique, Nancy, France, for
providing seed of Withania riebeckii, Hardy Eshbaugh for seed
of Capsicum annuum var. aviculare and Peter H. Raven for as-
sistance with literature. Particular thanks are extended to C.
E. Freeman, University of Texas, El Paso, who provided sugar
analyses of nectars, and to Suzanne Eder, Southern Illinois Uni-
versity-Edwardsville for microtechnique. The authors are grate-
ful for detailed reviews of the manuscript by Thomas G.
Lammers and Charlotte Taylor.
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William G. D’Arcy1Richard C. Keating1
William G. D’Arcy 1999 12 16
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... He recognised three sections, comprising sect. Tubocapsicum (today, a valid genus with two species, D'Arcy et al. 2001), the monotypic sect. Pseudocapsicum Hunz. ...
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Shrubs or perennial to annual herbs, rarely trees, rosette-forming or ephemerals, sometimes with tuberous or gemmiferous roots, or with tubers or stolons; stems occasionally with heteroblastic growth or with cauline spines; plants glabrous, frequently viscose or slightly or densely wooly-tomentose, with a diverse array of non-glandular and glandular trichomes. Leaves alternate, often in pairs, sometimes becoming opposite in the inflorescence, usually simple, entire, infrequently pinnatifid to deeply dissected or compound, exstipulate. Flowers perfect, rarely functionally unisexual in dioecious or andromonoecious plants, sessile to mostly pedicellate, fragrant or not, solitary or more commonly in axillary, extra-axillary, or terminal multi-flowered lax panicles, cymes or fascicles. Perianth (4)5(6)-merous; calyx actinomorphic, rarely zygomorphic, undivided or slightly to much divided, usually persistent and variously accrescent; corolla actinomorphic or zygomorphic, rotate, stellate, tubular, infundibuliform or hypocrateriform, exceptionally papilionaceous, tube glabrous or hairy inside, lobes longer or shorter than the tube, similar or dissimilar in size, aestivation valvate, valvate-induplicate, valvate-plicate, valvate-conduplicate, valvate-supervolute, cochlear, cochlear-conduplicate, cochlear-plicate, contorted-induplicate, contorted-conduplicate, contorted-plicate, quincuncial, or reciprocative; androecium included or exserted, 5-merous, 6-merous (Goetzea), 4-merous (then stamens equal in Nothocestrum, otherwise didynamous: 4 fertile or 2 fertile stamens and 2 staminodes; sometimes also with a fifth staminode) or with 2 fertile mobile stamens in lateral or dorsal position and 3 staminodes with vestigial or sometimes without anthers, filaments straight or declinate, inserted at different heights on the corolla tube, distinct or connate in a basal ring fused to the corolla, filament base appendages adnate to petal tube (“stapet”) absent, inconspicuous or conspicuous (with or generally without basal auricles), anthers dorsifixed, basifixed, dorsi-basifixed, or ventrifixed, extrorse, introrse or latrorse, frequently connivent (in Solanum sect. Lycopersicon with sterile apices and joined in a column), thecae generally non confluent apically, equal or unequal, dehiscence longitudinal or by terminal pores or slits, connective inconspicuous, wide, forming a dorsal layer of uniform and slight thickness, or thick with an emerging hump; gynoecium with oblique orientation (except Nicandra), usually bicarpellate, 3–5-carpellate in Jaborosa, Nicandra, Trianaea, and up to 30-carpellate in Nolana, ovary generally bilocular, sometimes 4-locular due to false septa, superior or sometimes partly inferior, glabrous or with trichomes or prickles; style heteromorphic or homomorphic, straight or declinate, terminal or subterminal, stigma variously shaped, usually wet and papillate (papillae rarely lacking); nectary absent, cryptic, or evident, then annular, invaginated or with 2 prominent lobes. Fruit generally a many-seeded berry or a septifragal, septicidal or septicidal-loculicidal capsule, rarely a pyxidium, drupe, diclesium or schizocarp. Seeds 1 to ca. 5000, compressed, then discoid, lenticular, reniform, irregularly ovoid or not compressed, then generally angular, cuboidal, ovoid, prismatic, polyhedric, subspherical, reniform; embryo straight to coiled, U-shaped only in Duckeodendron, cotyledons incumbent or oblique, less frequently accumbent, usually equal, slightly longer or shorter (up to 12 times shorter in the Australian endemic genera) and as wide or rarely broader than rest of embryo; endosperm rarely absent, generally copious, storing mainly oil and starch, with cellular type of endosperm formation (nuclear only in Schizanthus).
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