Systematics of the Guadua angustifolia Complex (Poaceae: Bambusoideae)

33 Pages·1992·20 MB·

by  Stephen M Young

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740 Annals of the
Garden
Missouri Botanical
economy and
the genus. These included the presence of alate to the local to the flora of the type
palea nondistichous inflorescences, three- locality" (McClure, 1957: 203). Kunth avoided
keels,
parted lower and a well-de- the use of a tautonym by changing B. guadua
styles, sterile florets, to
However,
fined geographical distribution. these ifolia
morphological characteristics are not completely la
life
consistent because they are known to appear in narrow-leaved characteristic has subsequently been
from both hemispheres. This Hackel found many species of the subgenus, and use
species led in its
Guadua
Guadua Bam- angus-
(1887) include as a subgenus of as a diagnostic feature limited.
to is
name
busa. Munro's (1868) monograph was influential ///b/Za the most often found in the literature
is
in the acceptance of Guadua as a genus, even and has been used in recent years (Burkart, 1969;
though he expressed doubts as to validity. After Rosengurtt et al., 1970; Purseglove, 1972; Azeve-
its
Guadua 1977a,
1868, the group was generally treated as do Correa et 1977; Azzini et al., b;
al.,
&
vmtil McClure (1973), uncertain of generic sep- Soderstrom Londoiio, 1987), although B. gua-
its
aration, reduced to a subgenus of Bambusa. dua has frequently appeared (Hidalgo Lopez, 1974;
it
In their study of the genera of American bam- Turpe, 1975; Rincon Sepulveda, 1977; Hidalgo
&
Maecha Vega
boos with pseudospikelets, Soderstrom Londono Lopez, 1978; Smith et aL, 1981;
&
Guadua 1983; Soderholm, 1984).
(1987) restored to generic status based Echeverri Restrepo,
on number
a of distinctive features, a trian-
i.e.,
gular culm leaf in which the margins of the sheath GUADUA ACULEATA RUPR. EX FOURN.
and
blade are contiguous or almost presence of
so,
Bambusa
name aculea-
Ruprecht used the
first
thorns on the culms and branches, a distinctive
annotat
7°
band "
of short white hairs both above and below the
'
m; e' specimens from \t^he i1o84^ 1i-lioo4/-i zo^iMv^ievxiircaaini ji"o"u»r-
j v J f r -1
hrm
nodalI Ime, and a pal1 ea ot 1t1ext1ure with prom-
t Wilhelm
wuneini
r Gr^erman ^*^r.
ney i i i coliilector
of the little-known
r
ment, wm. gs emanati. ng irom
the keels. "^ ''
after
von Karwin (Karwinsky). Fournier (1877),
We
are following most recent treatment of
this
name
used the
Karwinsky's specimens,
seeing
name
the group and use the Guadua. Monographic m
polygamy
Guadua of
aculeata discussion
in a
studies of the genus and related groups be
will
ap-
description
no formal
pseudospikelets, but
needed
to further clarify generic
limits.
Mexi-
Fournier's
of
peared the publication
until
were
specimens list^
Seven
Plantas
cartas (1886).
GUADUA KUNTH
ANGUSTIFOLIA
an
Veracruz,
from
ed as syntypes, five Colipa,
&
Humboldt Bonpland Bam- unknown Mexican
(1806) described two from localities.
Ruprecht s
guadua
busa and B, latifolia based on specimens Fournier also described another of
m
Guadua
made 1799-1804 name
during their South American ''herbarium species" with the
po-
the
voyage. These were the of the more than 30 termedia. The name probably referred to
first
New
species of World bamboos of the genus Gua- branch which are interiningle
of the leaves,
sition
dua be injhis
to described. In addition to Latin and French with the spikelets. Leaves are also P^^'^^^
and
descriptions, illustrations observations on
local ifol
uses and ecology were provided for both species. ference between the original descriptions ol
•
tne
of
The guadua
came length
specific epithet from the com- termedia and aculeata the
G.
is
a
mon name
aculea^^
used by
the people of Colombia. This spikelets, described as short in G.
^
^h^
name
is still in use and is usually pronounced gua- more than 5.1 cm long in G. intermedia,
^^^
a
du-a Colombia and are
in gua-du-a in Ecuador. Poiret spikelets seen from the type collection
-,
*-i'-i«r. from tlios
(1808), using Humboldt's cm
information, also pub- 2.5 and indistineuishable
lone are
^
i-i_j
two
„.^«Qr^
!• appears
lished an extensive description of B, guadua and Guadua intermedia
of G. aculeata,
discussed economic
its uses. more without f^^^^^'^ggj.
times the literature
in
When
(1822a) Bambusa (Hemsley
transferred
oration of Fournier's description
^
as
included it
latifc Ca mus McClure (1973)
1913).
,
be synonym
of B. aculeata.
_ , u^
McClure synonyrruzed
type. guadua
later selected B. McClure
as the Another
species that
(mean-
"The t«emts
type, citing, internal evidence as to which der Bambusa aculeata Guadua
is
[Kunth] distingi"^
ing which he was not able to
thornless),
mind
consists of the extensive
documentation by from of "^^"^^^^^^^^^^y
B. aculeata on the basis
Humboldt and Bonpland
63 (McClure,
(I.e. et seq) of B. their respective type collections
sp^
presumed
The
ifolia) in relation only between these
diflferences
Volume Number 4 Young &
79, Judd 741
Systematics Guadua
of angustifolia
1
f
upon
cies, reading the original descriptions, are solved specific taxon, and only recently Quarin
&
their thominess and spikelet size B. aculeata (1977), Soderstrom (1981), Nicora Rugolo de
(i.e.,
&
with seven florets per spikelet and thorns, and G. Agrasar (1987) and Londoiio Peterson (1992)
name
inermis with eight to nine florets per spikelet and chose to resurrect the B, chacoensis. Only
A
thomle thornless form of the similar species, two published photographs, both long-distance
s),
G. amplexifolia Presl, also occurs sympatrically views, are known to exist of B. chacoensis (Short,
with G. aculeata in southern Mexico and further 1975).
confuses assessment of taxonomic of G.
affinities
inermis. (This problem discussed in more detail PAST COMPARISONS
is
under
interspecific relationships.) Unfortunately,
These three groups of bamboos have never been
culm were
leaves neither nor
collected, illustrated,
com-
separated a key or adequately
satisfactorily in
\ described for these species when they were
first
pared. Guadua angustifolia has been separatt^l
cannot
Camus
by
from G, aculeata only once, (1913),
taxonomic
disposition of these names. Hitchcock
but the distinction was based on the presence (G.
(1913) Bamhusa
transferred aculeata and
G. to
aculeata) or absence (G. angustifolia) of thorns.
included Mexican
it in his listing of the grasses in
now known two bamboos show
that these
It is
the U.S. He
National Herbarium. mistakenly gave
various degrees of thorn development.
credit to Ruprecht an
for describing the species,
McClure (1955) included detailed descriptions
I error that was carried over McClure's (1973)
to
if
^ork. The Bamhusa
correct genus
citation in the
I
yfG
B. aculeata
IS (Rupr. ex Fourn.) Hitchcock.
He
same included two forms of G.
the key.
in
bam-
angustifolia in a separate key to introduced
bambusa
chacoensis
n. rojas
but none of the characteristics coincided with
boos,
The large bamboos of the subtropical regions of those included in the key to native species con-
Argentina
and Paraguay have rarely been de- taining G. aculeata,
been
scribed
in the literature. Nicolas Rojas Acosta, an
Guadua
from alia
unor ered different if
ods and fragmentary has never been separated from in a
descriptions, described these therefore it
bamboos seem be aware of G.
unde Rojas Acosta did not to
^_ key.
...„
,
(Rojas Acosta, and made no attempt to compare B.
1918). His description was so vague angustifolia
&
as to encompass many now chacoensis with this species. Londono Peterson
of the species included
m
and
the genus, but and common (1992) compare G. angustifolia, G. trinii G.
his locahty data the
name
he and conclude that G. cha-
used, "tacuara," from the chacoensis in a table
distinguish
it
more
other bamboos that grow range Guadua coensis a separate species closely related
in is
its (i.e.,
Paraguayana
Doell ex Mart, and paniculata
G.
Munro, information from the entire range
*1a picanilla"; and G. (Nees) Rupr. do not include
trinii
and
and exclude important char-
G. tagoara
(Nees) Kunth, Un- of G. angustifolia
'Hacuarazii").
lortunatelv. hp rf^a^iy^A^A ^^t,,*.^ Uo^if ^^ u^c x^^w. acters such as size ratio of culm leaf blade to the
barium
and culm
generally did not collect specimens entire leaf.
(Krapovickas,
1970). Consequently, he did not col-
^J a specunen
of bamboo
this Methods
and
Materials
type
specimen.
In 1931 Bam- This investigation involved field studies, tradi-
this species appeared a key as
in
bu methods herbarium spec-
taxonomic utilizing
tional
similarity
analyses of the
and multivariate statistical
mboos Guadua imens,
to
angustifc
di Guadua angustifolia complex-
if
Parod
dis
HERBARIUM STUDIES
'^uent
literature these bamboos have bee
microscope was used
Wild dissecting
1943;
ifc
rod amme pecunens
Young
B
reprod
urkari, specimens
for
1969;
Rosengurtt 1970) or B.
et ah,
g^fidua n ocular
Mc-
(Roseveare,
1948; Turpe, 1975).
microm-
"*"€ (1973)
included B. chacoensis as an unre
.
742
Annals
of the
Garden
Missouri Botanical
Table
1. Foliage leaf characters used in principal Table 2. Foliage and culm leaf characters included
=
PCA
components analysis. * omitted from analysis of culm in second with foliage leaf characters.
and
foliage leaf data.
Thorn development
1.
Leaves per complement Culm
1. 2. leaf sheath length
mm
>
2. Position on sheath of hairs sized 0.1 3. Culm leaf sheath base width
< mm*
3. Position on sheath of hairs sized 0.09 4. Culm leaf sheath ligule width
mm
4. Density on sheath of hairs sized >: 0.1 5. Culm leaf blade length
5. Degree of stramineous coloration in sheath 6. Culm leaf blade to culm leaf ratio
6. Degree of brown coloration in sheath 7. Culm leaf sheath vesture
Degree
7. of purple coloration in sheath 8. Culm leaf blade vesture
Degree
8. of green coloration in sheath 9. Contact of ligule with culm leaf margin
Number
9. of oral setae Density of culm
0. leaf ligule cilia
1
10. Length of oral setae Length culm
11. of leaf ligule cilia
Degree
1 1 of stramineous coloration in oral setae
Degree
12. of white coloration in oral setae
Degree brown characters) are analyzed in four separate catego-
13. of coloration in oral setae
14. Degree of purple coloration in oral setae ries, two vegetative and two reproductive.
15. Sheath auricle length The vegetative data includes 42 char-
set
first
Leaf
16. auricle width measured from leaves of 105
acters the foliage
17. Collar length
Table
specimens. These characters are listed in I.
18. Collar
flaring
The second vegetative data set includes selected
19. Ligule height
from
branch leaf data plus additional characters
1 1
< mm*
20. Position on ligule of hairs sized 0.09
These
culm leaves and branches of 65 specimens.
mm
>
21. Position on ligule of hairs sized 0.1
Another
Table
additional characters are listed in 2.
22. Pulvinus
height
24 were measured but omit-
vegetative characters
Degree
23. of stramineous coloration on pulvinus
inadequate her-
from because of
24. Degree of brown coloration on pulvinus ted the analyses
or
25. Degree of green coloration on pulvinus barium material, distortion in pressed material,
26. Degree
of purple coloration on pulvinus* extreme
their variability.
27. Pulvinus vesture 33 char-
The reproductive data set includes
first
mm
<
28. Position on adaxial petiole of hairs sized 0.09 features
acters measured by examining the external
mm
29. Position on adaxial petiole of hairs sized >: 0.1 _r -.n _^:i „i„^„ ^r 4a9q scpnepcriimmeennss^.
of inflorescences a_ n_ d_i spikelets of
mm
<
30. Density of adaxial petiole hairs sized 0.09
The second
These Table 3.
characters are in
listed
mm
31. Density of adaxial
petiole hairs sized >: 0.1
features
aU external
^^P^^ductive category includes
< mm
32. on
Position abaxial petiole of hairs sized 0.09
> mm* seven mternal characters from dissected p
mm* P'^s
33. oonn
PPoossiittiioonn aabbaaxxiiaall poeettiioollee ooff hhaaiirrss ssiizzeedd >: 0.1
1
character
< mm of 30 specimens. These additional
34. Density of abaxial petiole hairs sized 0.09 lets
were
33 characters
35. Density of abaxial petiole hairs sized >: 0.1 mm* are Hsted in Table 4. Another
because
36. Blade width measured and omitted from the analyses
37. Blade length
they proved to be too variable.
38. Blade width to length ratio* A reproductive char-
52 and
of vegetative
total
1
39. Blade adaxial hair distribution stu y-
course ol
acters were measured during the
40. Blade adaxial hair density
The sti^fJ^^^/
absence of any previous detailed
41. Blade abaxial hair distribution num
large
group the use of a
this necessitated
42. Blade
abaxial hair density
adequate
y
understand
of characters in order to
group.
the pattern of variation of the
Compo-
The data were analyzed by Principal
Each CLUSIA^
specimen usuaUy
eter.
consisted of several nents Analysis (PCA) using the program
ordinaUon
sheets-
(Wishart, 1975); see discussion of this
present
are
technique Wiley Results
(1981).
in
NUMERICAL pap
TECHNIQUES
of this
in the taxonomic treatment section
The
fraementarv and
Species Concept
boo herbarium
specimens
prevented
the simulta-
neous
gathering of delimiling
detailed vegetative and of
repro-
In the Bambusoideae the challenge
ductive data from absence of
but a few herbarium
all specimens, and by the
ranking complicated
taxa
is
In order to mclude as many specimens A reproductiv
as
possible, complete herbarium specimens.
years
data (consKstmg
of continuously many
varying
or "count'*
system after
that produces flowers only
Number
4 Young &
Volume Judd
79, 743
1992 Systematics Guadua
of
angustifolia
Table 3. External reproductive characters used in Table 4. Interior spikelet characters used in prin-
components
principal analysis. components
cipal analysis,
Number
of lateral leaves in inflorescence Lowest
1. rachilla section length
1.
Number
2. of terminal leaves in inflorescence 2. Lowest rachilla section width
3. Spikelets per cluster Lowermost palea length
3.
4. Spikelet length 4. Lowermost palea width between keels
Spikelet width
5. 5. Style length
6. Spikelet stature (erect to arcuate) 6. Style width
Number
gemmiparous
7. of bracts
7. Style hair length
Uppermost gemmiparous
8. bract length
Uppermost gemmiparous
9. bract width
10. Degree of stramineous coloration bract
in
with that of variety as proposed by Raven (1974).
Degree brown
11. of coloration bract
in
Both categories have been used sparingly by bam-
Degree
12. of purple coloration bract
in
boo taxonomists (Munro, 868; Camus, 1913;
1
Degree
13. of green coloration bract
in
Holttum, 1958; McClure, 1973), but study
Number in this
14. of veins on bract
15. Density of hairs at base of bract subspecies will be used to delimit populations that
and
16. Density of hairs at center of bract are geographically isolated exhibit variation,
17. Density of hairs at apex of bract which results in slight morphological gaps of a few
18. Density of hairs on edges of bract vegetative and/or reproductive characters. Pop-
Number
19.
of
florets ulations that vary in a few vegetative and/or re-
20. Lowermost lemma
length
productive characters and show some geographic
21. Lowermost lemma
width
continuity but no consistent morphological gaps are
22. Lowermost lemma mucro
length
geographic
referred to as ''incipient races.**
23.
lemma
24. Degree of brown on lemma
coloration
and
Interspecific Relationships
25. Degree
of purple on lemma
coloration
Definition of the Guadua angusiifolia Complex
26. Degree
of green coloration on lemma
Number
27.
of veins on lemma
Guadua
The genus in need of revision, since
is
28.
Density lemma
of short hairs at base of
many of the species have been inadequately de-
29. Density of short hairs at center of lemma
However,
scribed and collected. the species of the
30. Density of lemma
short hairs at apex of
complex from
G. angustifolia are clearly separable
31.
Density of short hairs on edges of lemma
The
32. Density the remaining taxa. combination of characters
of long lemma
hairs at base of
33. Density of long lemma that define the G. angustifolia complex are as
hairs at center of
(7-)10-20(-
culms of large stature
follows: (i.e.,
m
30) erect with arched tops; internode walls
tall),
of medium culm
vegetative growth of thickness; leaves large, abaxially
drastically reduces the op-
Portunity pubescent, and usually without auricles; foliage leaf
to observe, and experiment with
collect,
reproductive sheath glabrous or pubescent toward midrib of low-
McClure 973)
structures, a
situation
1
(
"•tened sometimes approaching the apex but never
to the system of "Fungi Imperfecti" de- er half,
veloped
by ibo
mycologists. These problems have lim-
,
cm
our and 1-4 wide, surfaces glabrous or only
knowledge long
Jjed of the processes involved in
bamboo
glabrous along midrib
pubescent, petioles
speciation and have kept bamboo taxon- sparsely
o^y 2-
from Synflorescence consisting of
advancing much of abaxial surface.
beyond the alpha stage.
on
pseudospikelets delicate.
multiflowered
5
'boo have been based
classifications
cm
2-9
strictly upon branching twigs; main florescence long
overaU simUarity and the presence of
mm
"""^ 3-14;
3-8 surface of bracts
g^ps
i' n observed and wide; florets
' than
morphological patterns rather
I
lemmas glabrous or with sparse hyaline hairs
pothesized and
evolutionary changes or biological in-
-
and edges, rarely densely pubes-
^^ractions. Whil base, center,
at
margins glabrous; palea
^traints base and center,
of time and above cent at
the problems referred to
beyond
^ve pubescent margins exserted the
limited keels with
the
present study to a simil
^
Proach.
lemmas; lodicules 3.
grow
genus do not taller than
Most plants in the
delimited
Guadua
angustifolia usually
10 m, while plants of
numbe
m
and 20 and are often
etativ between taller,
grow
to 1
I
Th.
^^^y 30 m. otbcr large species
reaching
of sometimes
subspeciXwrn
b^c^ons^^^^^^^
%
744
Annals of the
Garden
Missouri Botanical
of the genus are few and are easily separated from termlne the precise taxonomic relationship of
the
by and "hollow culm
G. angustifolia several vegetative repro-
ifoli
and
ductive characters. In Tables 5 a detailed
6,
made among
comparison
the species that are
is
PhytogeogRAPHY
Guadua
sympatric with angustifolia.
Guadua Guadua
angut
Although the species sympatric with
ifc
angustifolia are easily separated from this species range of any species in Guadua. Its northernmost
on vegetative and floral characters, the scarcity of localities are found in the state of Veracruz, Mex-
Mexico and and southernmost are found
G. angustifolia subsp. angustifolia in ico, its localities in
Central America (formerly known as G. aculeata) northeastern Argentina and southeastern Para-
Two
has caused problems in the past. forms of G.
^fi
amplexifolia are more common than G. angus- species that are known from both Central and South
^i/o/m Mexico and Central America. As a America; the others are G. paniculata and G.
in result,
G. amplexifolia has often been misidentified as G. plexifc
such
aculeata. This problem developed because of the Speculation on the factors resulting in ex-
lack of detailed vegetative descriptions and a veg- tensive distributions difficult because of the pau-
is
etative key separating the two species. Both G. city of information on the mechanisms of dispersal
amplexifolia and G. angustifolia are very thorny, of these large bamboos. Discontinuities in flowering
and no
and early collections were given the name G. acu- periods can be very long, up to 35 years,
leata. Once these collections were misidentified the studies have been made to ascertain the vectors
was seed-dispersal dis-
error carried over to the determinations of responsible for seed dispersal or
must
later specimens. Another problem concerns the tances of these bamboos. Vegetative dispersal
range;
certainly be a factor in increasing their
iplexifc
The form have been done show the ease with
ifolia. thornless of G. angustifc studies that
named when (McClure, IvoOj-
Mexico, originally G. merm/5, was described which culms root planted
nv-
grow along
from flowering material from Colipa, Veracruz. The These large bamboos frequently
m means
culms were described as 16 and unarmed, erbanks, and could serve as an efficient
floods
tall
downstream.
and the branch leaf sheaths and petioles were de- of vegetative dispersal, especially
or
continents
between
scribed as glabrous, characteristics of G. an- Long-distance dispersal
all
when culm sectio
gustifolia. There are some clumps of G. angus- islands could possibly occur
No na\
studies
tifolia in other parts of range (mainly the are carried by ocean currents.
its in
ex
prolonged
province of Esmeraldas, Ecuador) that have few tested the viability of culms after
o
has
When
or no thorns on the upper branches. branch- posure seawater, but the author
to first
ol
on kilometer
es the lower third of the culm, which are thorny, served clumps growing within a
Raven
fail to develop, the culm could be considered thorn- ocean and on islands within tidal bays.
Guadua found
bamboos
less. inermis likely has this same growth od (1974) sueeested that
times
Neogene
form. Unfortunately, there no complete vege- of Panama probably arrived in
is
tative specimen of G. inermis from Mexico even more from South America. It is ^
and, recently
nav
Guadua
because occurrence
its is so restricted, no living teresting that only three species of
may more intrigu-
plants remain. and even
a bicontinental distribution,
native
bamboos are
no
ing the fact that large
ifolia, is
many
which occurs in areas of Central America, West
This
has not been described in the literature and thus a).
ommon
has been
often
misidentified as G. inermis.
It
is
means
of dispersal.
ifc m
j r
fro^
found
culm Even though Guadua angustifolia is
m
gaps
and
leaves, foliage leaves (see Table These Mexico are large
Argentina, there
5), to
Amer-^
been Central
Throughout
2-4).
distribution
(Figs.
McClure
be
separated from common
the thorny ica not very and, as \ ^
is
it
m
from
suggested, may have been extirpated
it
but thick-walled culms. Thus^we have chosen cutting of its
to of former range by selective "
its
WhUe
wrote,
he
^'hoUow cuhn perior culms. In personal notes
his
spoK
who
n^ V
Moore,
be »^ tU^ R^e..
r^r.^A\,^^ i,.*^
y^^
as
reduced numbers. More many
in
study needed from
is to de- personal experience covering
-i
CD
Table
5. folia
i
3
Cundua
Guadua
angustifolia amplexifolia Guadua amplexifolia
CD
subsp. angustifolia culm" "hollow culm" Guadua Guadua
''solid longifolia paniculata
m
Culms 10 30 10- 8-10 m 6-10 8-10 m 6-10 m m
erect; tall, erect; tall, erect; tall, clambering; to 15 long, weakly erect; 8-9
tall»
20 medium cm cm
in diam.; diam.; solid with diam.; thick-walled to 5 cm diam.; thick- 5-7 cm diam.;
thick-
wall thickness pithy center wa
lied walled
Culm
brown
leaves pubescent; decidu- brown pubescent; brown
tardily pubescent; tardily slightly white pubescent; yellow-green with purple
ous; blade about Va or deciduous to persistent deciduous to persistent promptly deciduous;
horizontal stripes, pu-
less as long as leaf, and rotting; blade VS to and rotting; blade V^ to blade about Vi as long bescent; sheath
tardily
persistent; auricles ab- Vi as long as leaf, per- 14 as long as leaf, per- as leaf, base much low- deciduous; blade
Vi to
sent sistent; auricles present sistent; large auricles er on one
side; auricles Vi as long as
leaf, ca-
absent
to present absent ducous from
sheath;
auricles absent
Thorns
1-5
present to rarely absent; present; at node; absent to very sparse; few present; or rarely 2
1 at present; usually 3
at
i-5
at the node; short short to long at node if present; node, pointing down- node;
fairly long, slen-
C/)
O
to long
short ward
der and very sharp
CO
Branch leaves blades medium medium
width; blades to wide; blades wide; sheaths usu- blades very narrow;
blades narrow, pendant; CD
CO
sheaths glabrous or pu- sheaths glabrous
to to- ally glabrous; abaxial sheaths glabrous
to sheaths pubescent;
ab-
bescent along midrib of tally pubescent, usually petiole glabrous; auri- rarely pubescent; abaxi- axial petiole pubes- o
lower half; abaxial peti- on edges of lower or cles preseru al petiole glabrous to cent; auricles absent a
Q
ole glabrous; auricles upper half; abaxial peti- pubescent;
auricles
absent to rarely present ole glabrous; auricles present absent
to
absent present
to
Spikelet short lo long, narrow, flat- long and wide, rarely same as "solid culm" very long and narrow, and
short narrow,
flat-
1-3
tenod; per cluster; short, round; usually 3 round; 1-3
per cluster; tened; usually 3 or
glabrous more
lo sliglitly pu* or per cluster; glabrous more
per
cluster; pu-
t
bescent glabrous to rarely pu-
bescent
bescent
^
S
en
Tabi E 6, ifolia
Guadua
angustifolia
Guadua paraguayana Guadua Guadua tagoara Guadua paniculata
subsp. chacoensls trinii
10-15 m 8-10 cm 8-10 m 2-4 cm 8-10 m 3-5 cm 8-10 m 6-8 cm 8-9 m 5-7 cm
Culms
tall, tall, tall, tall, tall,
diam. diam. diam. diam.
diain.
Culm leaves brown pubescent; blade !4 brown pubescent; blade VJ brown pubescent; blade V2 brown pubescent; blades yellow-green with purple
much
or less as long as leaf, as long as leaf, persis as long as leaf, persis- as long as leaf, horizontal stripes, pu-
Vt
persistent; auricles ab- tent; auricles present tent; auricles absent wider than long, persis- bescent; blade V4 to Vi
sent tent; auricles absent as long as leaf, cadu-
cous from sheath; au-
absent
ricles
>
1
Branch leaves blades medium to narrow; blades medium to narrow; blades medium to narrow; blades wide; sheaths pu- blades narrow, pendant;
en
sheaths glabrous; abaxi- sheath with hairs on sheaths glabrous to pu- bescent on one side of sheaths pubescent; ab- CO
o
al petiole glabrous one or both sides of bescent at apex; abaxi- apex; abaxial petiole axial petiole pubescent
O
apex; abaxial petiole al petiole glabrous to glabrous
o
pubescent on one or pubescent
CD
0)
both
sides
o
Spikelets long and wide, glabrous to long to short and narrow, short and narrow, gla- short and narrow, gla- short and narrow, pubes-
slightly scabrous; palea pubescent; palea wings brous; palea wings brous; palea wings cent; palea wings not
wings
exserted, ciliate usually not exserted, broadly exserted, gla- broadly exserted, not exserted,
ciliate
brous minutely
ciliate to cili- ciliate
CD
3
ate
Number
Volume 4 Young &
79, Judd 747
1992 Guadua
Systematics
of angustifolia
I
Figure
2. native
ifolia
=
plants;
triangle
possible cultivated plant.
a
consultant on land and savanna (Harling, 1979) as
ciduous
utilization in several countries forest,
of
Central America, and edaphic conditions
thousands of acres of land long as water plentiful,
is
originally
occupied by more This
or pure stands of are satisfactory
less
^mboo
more
[G. amplexifolia and aculeata] have made the species resistant to past climatic
G.
heen
Many America where
cleared areas of Central
in recent decades for agricultural changes. it
purposes.'' Unfortunately, bamboo stands were could possibly occur have yet to be explored bo-
synonymous ^ J^^^ * M w M V V
_ Ink rik bfl bean
with "good banana
land."
rast
climatic changes may have decreased the be
distribution
described
of the habitats available, as Toledo ^„
.
. -
U982)
described in the changes undergone by the of the conspicuous* features of the landscape in the
' Guatemala and Honduras, being
* Within 10,000 regions of
the coastal
' last
times abundant swamps and along rivers at low ele-
extent of the in
i
tropical
forest must have forming extensive groves.
decreased considerably, vations, often
iboo bet
ere
there may specimens
are very few have
collections,
f^en Am-
eliminated Guadua Guyana. the
altogether. angustifolia Venezuela and Its distribution in
»
conspicuously known. The eastern
absent most of Nicaragua and azon basin remains very poorly
in
of many
ali Costa Rica, and even though occurs populations can be found along of the large
in
it
how
South Ecuadorian Oriente, but just far
of the
rivers
nown
bordering
'/ stifc
Colomb
ted
and orrurring widely
be^ found lowland taxonomic section) in
in rainforest, (see
ower
montane Subspe-
2-4).
rainforest, semideciduous forest, de^ separated gf»ngraphic ranpf:s (Fips.
748
Annals
of the
Missouri Botanical Garden
SUBi'^A
^^"'^",^"*'°" **f ^^ste^n (dots) and eastern Guadua angustifoU
(dots in circles) populations of
^*^V.^/,^- =
angustifolia. Triangle
possible
cultivated
plant.
ifc
bamboo-dwelling
basin of Peru and
subspecies reported.
chacoensis has 483)
its southeastern Peru, Parker (1982:
northernmost
distribution
in eastern Paraguay reiatiy^
Most
close
"It interesting to note that
IS
of the specimens
of G. angustifoU
Flamm'
Hemitriccus
flammulatus, the
coensis have been
collected along Slate-
th. the
„.,^ my-Tyrant and Sporophila shistacea
Paraguay
rivers, but not known ocnruyjo
it is
the sub" Drymophila
if
Seedeater
(i.e.,
speces
extends up
these nvers
„^A ^r^nroDhilnP*
into BrazU and ._„x_j Sporop
Drab-breasted Pr.ygmy-TTy- rant. and
if