Reproductive Morphology, Anatomy, and Relationships of Ticodendron

8 Pages·1991·7 MB·English

by  Hiroshi Tobe

REPRODUCTIVE
Hiroshi Tobe^
MORPHOLOGY,
ANATOMY, AND
RELATIONSHIPS OF
ON
TICODENDR
Abstract
Ticodendron predominantly dioecious. Both male and female inflorescences bear some cymules
is (or partial
on primary Male
inflorescences) their axes. partial inflorescences are borne verticillately in clusters of three, each
8-10
being one- to three-flowered and subtended by a single bract. Staminate flowers have or more stamens and lack
a perianth. Female partial inflorescences are solitary. Each of them always one-flowered, although subtended
is it is
by three (one primary and two secondary) bracts, and produces some scales in each axil of the secondary bracts.
it
In pistiUate flowers, concentric bundles are prevalent in the periphery of the pistil wafl; the ovary inferior, bicarpellate,
is
and four-locular with one ovule in each locule; the placentation axile; the ovules are apical, pendulous, hemitropous,
is
20-30
unitegmic, and crassinucellate; the integument is cells thick and vascularized by about 20 longitudinally parallel
vascular bimdles; and the mature seeds have a two-cell-layered endosperm and seed coats composed of thick-walled
integumentary cell layers and vascular tissues. In reproductive morphology Ticodendron from any other
distinct
is
On
family but best placed in Fagales. the basis of evidence from reproductive morphology and from other sources
is
common
it is suggested that Ticodendron appears to have diverged early from a ancestor with Betulaceae or Fagaceae.
T &
Ticodendron^ comprising only incognita, 1977) as described in Tobe Raven (1984). Sec-
6-
which occurs from northern Nicaragua through tions cut at 10 fim thickness were stained with
Panama,
Costa Rica to central has long been a Heidenhain's hematoxylin, safranin, and fastgreen
mystery plant because of uncertain identity (Go- FCF.
its
& &
Gomez Hammel
mez-Laurito 1989; Bur-
P.,
The
ger, 1991). present paper provides a detailed
OBSERVATIONS
morphology and anatomy
description of of
flora!
Ticodendron mostly Male and
dioecious.
fe-
Ticodendron few remarks on em- is
including a
its
How-
inflorescences both appear be
"^^'^ to spikes,
upon
bryology, in order to shed hght the position
on
ever, the basis of anatomical observations
dis-
and
of the genus.
affinity
cussed later, has been found that the "spike"
it is
composed some
of one- three-flowered cymules
to
Materials and Methods
To
borne on the primary axis. facilitate discussions
examined were Mon-
All materials collected at
comparing Ticodendron
in with other related taxa,
FAA;
teverde, Costa Rica, and fixed with examined more
a comprehensive term, "partial inflores-
were mature male {Haber
nearly flowers 7072),
111 cence" (instead of "cymule"), be used the
li^»/ will in
^ ^ #
^^>
young female buds and {Gomez-Laurit* -o
fruits s.n. 164-165,
following descriptions (see Abbe, 1974:
Haber
1986; 6840, 6868, 7286). Voucher
in
terminology).
fQj-
MO
specimens are deposited and CR.
at
For anatomical groups
study, single flowers, of
male and flowers
partial inflorescences
male and female flowers, and a piece of mature
seed were sectioned with a rotary microtome fol- Male partial inflorescences are borne verticil-
lowing standard paraffin methods. Before being sec- lately on the inflorescence axis in clusters of three
tioned, the materials were softened with a mixture (Fig. Each male partial inflorescence sub-
1). is
10%
of 10 parts of glycerol, 3 parts of Aerosol tended by a single primary bract, and composed
is
&
OT, and 90 parts of water (Schmid Turner, of 1-3 staminate flowers basically arranged in a
am
grateful to Jorge Gomez-Laurito, Barry Hammel, and William A. Haber for collection of fixed materials,
* I
Grayum Gomez
to Michael H. and Luis Diego for valuable information on Ticodendron and related taxa in earlier
stages of the work, and Peter H. Raven support various ways.
to for in
^ Department of Biology, College of Liberal Arts and Sciences, Kyoto University, Kyoto 606, Japan.
135-142.
Ann. Missouri Box. Card. 78: 1991.
136 Annals of the
Garden
Missouri Botanical
6D-H).
dichasium. Individual partial inflorescences (as well of flower (Figs. 5, This peripheral vascular
as individual staminate flowers in the case of three- system supplies the perianth (and abortive stamens,
Each
flowered partial inflorescences) are not readily dis- if present) and the styles (Fig. 6F-I). style
tinguished from each other, but their distinctness has three (occasionally four) vascular bundles (Fig.
revealed by vascular bundles. Those supplying
61).
is
each partial inflorescence or each staminate flower The central vascular system is composed of some
depart in separate groups from the vascular cyl- 20 small collateral bundles, which ascend as pla-
inder of the inflorescence axis. Staminate flowers cental bundles to supply the ovules (Fig. 6E, F).
8-10
have or more stamens and lack perianth and Ovule placentation axile.
is
have an
or rarely they abortive (see
pistil, pistil
OVULES
arrow.
Fig.
1).
The stamens are tetrasporangiate, and the an-
Although were
flowers collected simul-
pistillate
Anther comprise
thers dehisce longitudinally. walls
mature
taneously with nearly staminate ones, the
a fibrous endothecium and a persistent epidermis
were megaspore dyad
oldest ovules in the stage.
still
when
maturity. Pollen two-celled shed.
at is Probably more than several weeks pass between
and The
pollination fertilization. oldest ovules avail-
FEMALE PARTIAL INFLORESCENCES AND FLOWERS able indicate that the ovule apical, pendulous,
is
and hemitropous with the micropyle upward, and
Female and
partial inflorescences are solitary,
somewhat grows The
that twists as (Fig.
7).
it it
each always one-flowered, although sub-
is it is
By
ovule megaspore dyad
crassinucellate. the
is
tended by one primary and two opposite secondary
4-5
about above dyad
stage, parietal the
cells lie
may
These
bracts. three bracts fuse basally (Fig.
(Fig. 8). Obturators are lacking.
At each secondary some
6B). axil of the bracts,
The
ovule unitegmic (Fig. with the integ-
is 7),
10 small appendages or scales are produced, each
20-30
ument
about thick and with
cells tips
its
receiving a vascular bundle These
distinct (Fig.
2).
forming a long narrow micropyle. The integument
and
scales are horizontally scattered in the axil are
some 20
vascularized by longitudinally parallel
is
somewhat
attached
at different levels.
vascular bundles which are derived from
(Fig.
9),
composed
Pistillate flowers are only of a single
a raphe bundle via post-chalazal branching. The
its
inferior ovary with a small, inconspicuous perianth
integument each
longitudinally ribbed, con-
is rib
on ovary The
the top of the (Fig. flowers rarely
3).
taining a single vascular bundle (Figs. 9, 10). Outer
have abortive stamens on the inner side of the
epidermal of the integument are
cells radially elon-
perianth; the abortive stamens receive a distinct
gate and contain cytoplasm
rich (Fig, 10).
The
vascular bundle as do ordinary ones. ovary
is
two
bicarpellate with (rarely three) elongate sep-
MATURE
SEEDS
arate styles and four-locular, each locule containing
Only one of the four ovules the ovary develops
in
one ovule Styles are stigmatic throughout
(Fig. 4).
into a mature seed, thus each mature one-
A fruit is
with elongate and papillate epidermal nar-
cells.
seeded. The embryo of a mature seed and
oily
row adaxial sulcus present on styles throughout. is
is
massive with two straight cotyledons. The mature
Judging from the position of the two styles in re-
seed has a two-cell-layered endosperm
(Fig. 11).
lation to the inflorescence axis (see Fig. 6C, the
I),
As
the seed matures, only the outer tissue of
two
carpels are positioned tangentially, not radially.
the integument, including a vascular tissue and a
The anatomy
vascular of a female flower
floral
surrounding integumentary
while
tissue, persist,
presented Figure which diagrammatically
in 6,
is
the inner tissue completely degenerates. Thus, the
shows the vascular course in a longitudinal section
8-15
mature
seed coat only about thick
cells
is
6A) and
(Fig. in selected serial transverse sections
Those xylem
(Fig. 11). cells, except for cells of the
6B-I). At and above the which the
(Fig. level at
vascular tissue, are thick-walled and flattened with
scales are supplied with vascular bundles (Fig. 6B,
The
tanninlike contents. outer epidermal of
cells
C), the vascular cylinder, which composed of
is
the integument, which were elongate and
radially
many
discrete collateral bundles arranged in a ring,
contained rich cytoplasm ovular stages, do not
in
irregularly diverges into the peripheral vascular
show any mature
specialization in the seed coat.
system and the central vascular system (Fig. 6D,
E).
Discussion
The
peripheral vascular system about 5 or
is 1
4-10
The
16 aggregates of small bundles. Included are above-described inflorescence and
floral
a few inverted ones throughout the whole length structures of Ticodendron within the range of
fit
Volume Number Tobe 137
78,
1
Ticodendron Morphology, Anatomy,
1991
and
Relationships
®
f
®
—
Anatomy
Figures 1-5. of partial inflorescences and flowers of Ticodendron. Transverse section (TS) of a
1.
Arrow
primary male inflorescence axis showing three male, one-flowered partial inflorescences. indicates abortive
—
2. TS at the base of female partial inflorescence. Note that some scales are borne at the axil of the secondary
pistil.
— —
TS
bract. 3. Longitudinal section of female partial inflorescence, which is always one-flowered. 4. of female flower
—
TS
at level of pistil. 5. of concentric bundles prevalent in the periphery of the female flower. Abbreviations: br2,
mm,
secondary bract; single male flower; ov, ovule; pe, perianth; scale; sy, style. Scale bars equal 500 ^m,
f. s, 1
mm, mm,
and 100 ^m,
respectively.
1 1
138 Annals
of the
Garden
Missouri Botanical
Figure Diagrams anatomy
6. illustrating the vascular of female partial inflorescence and flower Ticodendron.
in
Median B-L
A. longitudinal section through the line presented in C. Transverse sections at levels marked b-i in A.
Abbreviations: ax, suppressed primary inflorescence axis; brl and br2, primary and secondary bracts; ov, ovule; pe,
perianth;
s, scale; sy, style.
Number Tobe 139
Volume
78,
1
Ticodendron Morphology, Anatomy,
1991
and
Relationships
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—
LS
Figures 7-11. Anatomy of ovules and seeds in Ticodendron.— 7. Longitudinal section (LS) of ovule. 8.
—
of ovule showing a dyad of megaspores in a nucellus. 9. Transverse section (TS) of ovule showing that the integument
ribbed and contains a vascular bundle in each rib.— 10. Enlargement of part of Figure 9, showing a structure of
is
em, embryo; endosperm;
integument.— TS mature seed showing seed coat structure. Abbreviations: en,
thick 11. of
vascular bundle. Scale bars
integument; mc, megaspore; mi, of micropyle; nu, nucellus; pr, parietal cell; vs,
tip
in,
100 ^m, 20 200 200 and 100 respectively.
equal fim, fim, fim, fim,
140 Annals of the
Garden
Missouri Botanical
Table 1. Comparisons of Ticodendron with related families in selected reproductive characters.
Fagaceae
Betulaceae Juglandaceae Myricaceae
Characters Ticodendron
(Fagales) (Fagales) (Juglandales) (Myricales)
Cupule
Absent Present Absent Absent Absent
Ovary
position Inferior Inferior Inferior Inferior Superior
Concentric bundles in Present Present Present
wall
pistil
Number
of carpels per 2
(2)3(-12) 2(3) 2(-4) 2
ovary
Number
of locules in 4 (2)3(-12)
2(3)
1 1
ovary
Number
of ovules 4 (4)6(-many) 2-4(-6)
in
1 1
ovary
Ovule
position Axile, pendu- Axile, pendu- Axile, pendulous Axile, erect Basal, erect
lous lous
Ovule curvature Hemitropous Anatropous Anatropous
Orthotropous Orthotropous
dumber
of integu-
1 (1)2 1(2)
1 1
ments
20-30 4-5
6-10
If uniteginic, thickness 4 3-7
cells cells cells thick cells cells thick
of integument
thick thick
thick
Vascular bundles
in Present Present Present Present Present
integument
Ribbed integument Present Absent ? Absent 9
*
•
Mature
seed coat Cells thick- Cells thick- ?
Cells thick-
*
walled
walled
walled
2-4
Thickness of endo- 2 cells thick Absent or 1 cells thick 1 cell thick 1 cell thick
sperm
thick
cell
References Vaughan
are: Netolitzky (1926), (1970), and Corner (1976) for families (seed coat anatomy);
' all
Langdon
(1939, 1947) and Poole (1952) for Fagaceae; Woipert (1910) for Betulaceae; Karsten (1902), Langdon
&
(1939), Meeuse Houthuesen (1964), Verhoog (1968), and Manning (1978) Juglandaceae; and Kershaw
for (1900),
Kuhn Yen
(1927), and (1950) Myricaceae.
for
shown
variation for these features in Fagales, Ju- Ticodendron agrees with Myricales in having a
glandales, and Myricales (see Abbe, 1974, for re- longitudinally ribbed integument.
view) and suggest closer affinity of Ticodendron Thus, the reproductive morphology and anato-
my
with these orders than with other Hamamelididae of Ticodendron indicates a close relationship
members Ham-
or of different subclasses. In Table 1, to Fagales, Juglandales, and Myricales in the
reproductive features of Ticodendron, Fagales, amelididae ''Hamamelidae" sensu Cronquist,
(or
Among
Juglandales, and Myricales are presented. 1981). Fagales, Juglandales, and Myri-
In general, Ticodendron agrees well with Fa- Ticodendron seems be more home
cales, to at
and They an
gales, Juglandales, Myricales. share within Fagales, which have diverse reproductive
inferior ovary (except Myricales with a superior morphology, rather than in Juglandales or Myri-
ovary), concentric bundles in floral vascular system cales, both of which are characterized by certain
(though uncertain in Betulaceae and Myricales), unique apomorphies. Juglandales and Myricales both
bicarpellate ovaries (though frequently tricarpellate characteristically have a unilocular bicarpellate
in Fagales and Juglandales), unitegmic ovules (com- ovary with a single orthotropous ovule, features
all
monly unknown
bitegmic in Fagales), vascular bundles in the that are Ticodendron and Fagales.
in
integument (in the outer integument only in Fagales Closer of Ticodendron with Fagales are
affinities
with bitegmic ovules), a similar mature seed coat further suggested by evidence from wood anatomy,
composed of numerous thick-walled cell layers and pollen morphology, and other data, as specified in
vascular tissues, and a thin endosperm in mature this volume.
Among may
Ticodendron
seed. further coincides with Fagales Betulaceae, Coryleae be the
closest
in having two (or more) ovules per carpel and axile allies of Ticodendron, since Coryleae, like Ticoden-
placentation, rather than a uniovulate ovary with dron, lack a perianth in the male flowers but possess
basal placentation as in Juglandales and Myricales. in the female flowers. Evidence from wood anat-
it
Volume Number Tobe
78, 141
1
1991 Ticodendron Morphology, Anatomy,
and
Relationships
omy
also suggests Coryleae for the closest affinity tinct from Ticodendron in having a basal, erect
of Ticodendron (Carlquist, 1991). Yet, in Ticoden- ovule and lacking a perianth in the pistillate flower.
dron the ovule hemitropous with a thicker in- Hjelmqvist (1948) suggests on the basis of com-
is
tegument (20-30 cells thick), whereas in Coryleae parison in the "cupule" structure that Balanops
anatropous and probably with a thinner integ- probably stands nearest Juglandales and Myricales.
it is
ument (e.g., four cells thick in Betula; Wolpert, Carlquist (1980) notes that the wood of Balanops
1910); Ticodendron has scales at the axils of the comparable with that of Hamamelidaceae. Za-
is
&
secondary bracts, which are unknown in Coryleae vada Dilcher (1986) indicate on the basis of
Ticodendron
(and in other Betulaceae); lacks similarity cluster analysis using pollen characters
all
the tertiary bracts that are positioned decussate to
^fagu
the secondary bracts and form by their fusion a the present level of our knowledge uncertain
it is
characteristic involucre or hull in Coryleae (Abbe, even whether Balanops related with Fagales
in
is
1935, 1938, 1974); in Ticodendron stamens are general. Balanops poorly known with respect
is to
many
not deeply divided as in Coryleae (and in other reproductive structures, and an intensive and
all
Betulaceae); and in Ticodendron female flowers overall investigation on the genus seems to be de-
are solitary, while in Coryleae two lateral flowers sired prior to critical comparisons with putative
of the dichasium usually develop (Abbe, 1935, and Ticodendron.
allies
1938, 1974). These differences suggest that Tico- In considering evolutionary relationships with
dendron unlikely to have direct relationships with Betulaceae and Fagaceae, the presence of the scales
is
Coryleae or other Betulaceae. in the secondary bract axils of female partial in-
may
Nothofagus the best candidate in comparing florescences of Ticodendron have a key
is role.
Fagaceae with Ticodendron, In fact, Nothofagus Although l-3-(or 7- or more-)flowered partial male
superficially similar to Ticodendron in having and female inflorescences are prevalent in Betu-
is
exclusively unitegmic ovules, occasionally solitary laceae and Fagaceae (see Abbe, 1935, 1938, 1947),
staminate flowers, partial inflorescences composed they are generally considered to have evolved from
of 1-3 flowers, and occasionally bicar- a more complex cymose inflorescence by reduction
pistillate
pellate pistillate flowers (for data on Nothofagus of branches of higher orders (see Abbe, 1938,
see Langdon, 1947; Poole, 1952; Abbe, 1974). 1974). Indeed, reduced outer branches are
sterile
&
Ticodendron, however, clearly distinguished from elaborated as cupular valves in Fagaceae (Fey
is
Nothofagus (and general from other Faga- Endress, 1983); furthermore, of presum-
in vestigials
all
ceae) in having hemitropous ovules with thicker ably reduced outer sterile branches appear as "su-
pernumerary
integument (only four or five cells thick in Noth- scales'* in male partial inflorescences
ofagus; Poole, 1952) and lacking characteristic of Chrysolepis chrysophylla (Fagaceae)
in
cupular valves, lacking a perianth in male flowers, (Hjelmqvist, 1948) or as blinding vascular bundles
and lacking the tertiary bracts in female partial in female partial inflorescences of Betula tenia
Those seem
inflorescences. dissimilarities to indi- (Betulaceae) (Abbe, 1935). Although uncertain
it is
cate that Ticodendron clearly stands out from Fa- whether the scales of Ticodendron are directly
gaceae as well. relevant to the scales of the cupule of Nothofagus
Balanopaceae (fia/arto/?5 only), which have been or not, their anatomical features are very similar
may
placed by Cronquist (1981, 1988) in Fagales, to each other in their cross sectional shape and in
also require comparisons with Ticodendron. Bal- receiving a distinct vascular supply (see Langdon,
anops shares with Ticodendron a female partial 1947, for cupule-scale anatomy in Nothofagus),
The
inflorescence comprising a single pistillate flower scales of the female partial inflorescence of
may
subtended by a "cupule,'' an ovary with locules, Ticodendron also represent residuals of re-
an anatropous, unitegmic ovule, and drupaceous duced outer branches of cymose
sterile inflores-
However, Balanops com-
the ''cupule" of cences.
fruits.
is
posed of numerous, spirally arranged, deltoid bracts. Regarding nonreproductive characters, Carl-
lacking an elevated axial structure characteristic quist (1991) showed that Ticodendron has primi-
of Fagaceae (Hjelmqvist, 1948). Its homology with tive wood compared with other Fagales; Feuer
the cupule of Fagaceae doubted (Cronquist, (1991) observed that pollen of Ticodendron had
is
Even
1981). the bracts of the ''cupule'' of Bal- plesiomorphic (tectate-granular) wall structure
if
anops are comparable with the scales of Ticoden- shared with Betulaceae, Nothofagus, Casuarinales,
&
t/ro«, the former encloses the whole of the pistillate and Urticales (see Zavada Dilcher, 1986); and
&
Hickey
flower, while the latter restricted only at the axils Taylor (1991) suggested, on the basis
is
Balanops
secondary
of the bracts. clearly dis- of leaf architectural analysis, that Ticodendron
is
142 Annals of the
Garden
Missouri Botanical
took a more basal position than other Fagales (and HjELMQVIST, H. 1948. Studies on the floral morphology
and phylogeny of the Amentiferae. Bot. Not., Suppl.
may
Brunelliaceae). Ticodendron be considered an
1-171.
2:
common
early derivative either from a ancestor of
^1902.
Karsten, g! Ueber Entwicklung der
die weibli-
Fagaceae before the reduced outer branches
sterile
chen Bliithen bei einiger Juglandaceen. Flora 90:
cymose were 316-333.
of the inflorescence elaborated as the
Kershaw, M. The and development
cupular valves, as in modern Fagaceae, or from a E. 1900. structure
Myrica Ann. (London) 23:
of the ovule o{ gale. Bot.
common
ancestor of Betulaceae before the reduced
353-362.
outer sterile branches were lost completely, as in
KuHN,
G. 1927. Beitrage zur Kenntnis der intrasemina-
modern For more
Betulaceae. critical discussion of len Leitbiindel bei den Angiospermen. Bot. Jahrb.
325-379.
61:
embryological Ticoden-
relationships, studies in
M
Langdon 1939. Ontogenetic and anatomical
L.
dron. Nothofagus. and Betulaceae, which are
still
and Fagaceae and
studies of the flower fruit of the
meager compared with those in other genera or
301-
Juglandaceae. Bot. Gaz. (Crawfordsville) 101:
are needed.
families,
327.
1947. The comparative morphology of the
.
Fagaceae. The genus Nothofagus. Bot. Gaz. (Craw-
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350-371.
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Abbe, E. C. 1935. Studies in the phylogeny of the W.
Manning, E. 1978. The classification within the
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1-67.
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1087.
1938. Studies in the phylogeny of the Betu- &
. Meeuse, a. D. Houthuesen. 1964. The
gy-
J. J.
laceae. 11. Extremes in the range of variation of floral noecium of Engelhardia spicata (Juglandaceae) and
and inflorescence morphology. Bot. Gaz. (Crawfords-
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its
431-469.
99:
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. Netolitzky, 1926. Anatomie der Angiospermen Sa-
F.
159-261.
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Carlquist, S. 1980. Anatomy and systematics of Bal-
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191-246.
anopaceae.
Allertonia 2:
Poole, A. L. 1952. The development of Nothofagus
Wood
1991. and bark anatomy of Ticoden- &
New
. seed. Trans. Proc. Roy. Soc. Zealand 80:
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207-212.
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Card. 78: &
ScHMiD, R. M. D. Turner. 1977. Contrad 70, an
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ical
An
Cronquist, A. 1981. Integrated System of Classi- &
An
Tobe, H. H. Raven. 1984. embryological
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York.
397-
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New
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