Table Of ContentA
OF COLICODENDRON
REVISION
(CAPPARACEAE)
Hugh
XavierCornejo
H.
litis
New
The
York Botanical Garden Department
of Botany
and
200th Una
Kazimiroff of 430
St. 'Ave. University Wisconsin,
New
0458-5
Bronx, York
126, U.S.A. Madison, Wisconsin 53706,
t
1
[email protected], [email protected]
[email protected]
During
our
studies of the neotropical
species of Capparis (Capparaceae), carried out to produce a generic
s.l.
realignment
extremely polymorphic became
of this genus,
clear that Colicodendron Mart, a well defined
it is
South American
genus of shrubs and by having
stellate trees, characterized a 1-seriate valvate calyx with
a
dentate or lobed nectary-dish and
or nectary-cup, thick-walled, indehiscent amphisarcous
fruits.
was by
Colicodendron established Martius two
(1839: 25), for Brazilian species: Colicodendron yco Mart.,
and
the generic
type, C. longifolium Mart, (the latter recently placed in Neocalyptrocalyx, as N. [Mart
longifolium
J
&
Cornejo H.H. Seeman
X.
[2008a]). Subsequently,
litis (1852: 78) correctly transferred Colicodendron:
to
Capparis scabrida Kunth, but also the unrelated Capparis Kunth, and
avicennifolia Capparis pulcherrima
Jacq.
nom.
(the latter as Colicodendron pulchellum Seem., reject.), as well as the quite unrelated glabrous or simple-
haired Capparis Kunth Two
subbiloba (=Cynophalla Turczaninow
years
flexuosa
Presl). (1854:
[L.] later,
J.
327-328)
described them synonyms
erratically three species in Colicodendron, all of of unrelated species of
Capparis a contribution that obfuscated any concept
s.l, clear of that genus.
was
Colicodendron subgenus
treated as a of Capparis L. (Eichler 1865: 272), and later as a section of
s.l.
&
Hoffmann
Capparis (Pax 1936: 181). In both mentioned infrageneric taxa, the members of Colicodendron
were mixed up
in heterogeneous assemblage with Mexican Kunth
a the Capparis
angustifolia (= Quadrella
&X.
[Kunth] H.H. and South American Kunth
angustifolia Cornejo), the and
litis C. crotonoides tweediana
C.
&
members
Eichler (both H.H.
of Capparicordis X. Cornejo, a recently described
neotropical
litis stellate
&
genus, Cornejo Most Hutchinson
2007). recently attempted
litis (1967: 309), in his generic realignment
made
of Capparis seven
additional, but inappropriate combinations produce
s.l, all to a poorly defined
Colicodendron.
In work, The
this a revision of Colicodendron presented. genus validated and amended, mor-
is
is
its
phological and
limits are defined, the relationships with the morphologically
closely related genera are
discussed.
Journal of the Botanical Research Institute of Texas 2(1)
76
&
emend. Cornejo H.H.
Colicodendron Mart., Flora 22, Bleibl. 1:25. 1839, X. litis. Type: Colicodendron yco Mart.
by trichomes throughout. Leaves simple, short-petiolate
Evergreen shrubs or covered stellate to echinate
trees,
subterminal and/or panicle,
whorled. Inflorescences terminal, axilar, a
opposite spirally or
to sessile, to
bracts filiform or
Colicodendron
raceme or racemose spike, rarely a solitary axillary flower (in yco)\ floral
subulate, deciduous. Flowers with calyx 1-seriate, valvate, ± cup-shaped, the 4 sepals free, ± equal, distinct
bud
from
margins and touching each other early until
and enclosing the corolla in bud, their tips
entirely
cup
forming hypantium, coated by 4-dentate nectary dish or nectary or
near anthesis, the base fused a a
at
bud, Stamens 14 the filaments
34obed nectary cup within. Petals imbricate or torsivus in sessile. to 60,
a 4,
on an expanded androgynophore,
arranged bud, anthesis exserted, inserted flat
geniculate or spirally in at
woody, thick-walled
pseudoamphisarcous
amphisarcous scabridum),
or (in C.
anthers basifixed. Fruits
mm
yellow-embryoned 1-2 cm, cotyledons 1-1.5 wide,
and bearing white- or seeds, ca.
multilocular, ca.
crustaceous.
convolute,
testa
from
Due pubescence, valvate calyx with the lobes covering the corolla early
the 1-seriate
to stellate
the
bud until near anthesis, and the presence of a nectary cup or nectary dish, Colicodendron is related to
&
However,
H.H. Cornejo. Colicodendron differs
described neotropical genus Capparicordis X.
recently litis
from by having oblong oblanceolate, pinnately nerved leaf blades (vs. cor-
the lanceolate, to
latter elliptic,
and subpalmatly-nerved 4-dentate nectary dish or
reniform suborbicular basally leaf blades); a
date or
to
number
appendages); higher
3-lobed nectary cup nectary cup with four filiform to triangular-filiform
a
(vs.
amphisarcous with an
of stamens, 14 to 60 4 to and larger indehiscent thick-walled fruits, inflexible
(vs. 8);
cm and
with thinly
woody 1-2 smaller dehiscent a soft flexible,
pericarp bearing larger seeds, ca. (vs. fruits,
mm,
5-7 by
dispersed
which and maturity, bearing smaller seeds, ca.
coriaceous pericarp, splits falls off at
South American genus Calanthea (DC.)
the
Cornejo 2002). Colicodendron also related to
birds, pers. obs., is
with Capparis pulcherrima
Miers, as was correctly stated by Martius in the original publication (p. 27, cf.
the type of Calanthea [DC] Miers). However, Calanthea mainly differs from Colicodendron by the very
Jacq.,
widely spaced
calyx with open aestivation, in which the not valvate linear-ligulate sepals are
distinctive
from very young bud; and by the seeds with barely folded,
from each exposing a valvate corolla the
other,
mm
&
4-7
wide (Cornejo 2008b).
very thick cotyledons,
litis
American-West Indian genus Qua-
Central
Colicodendron could be also related to the stellate to peltate,
hypanthium
(DC) However, Colicodendron differs from the latter by having flowers with a coated
Presl.
drella
J.
and
cup without hypanthium, but with a receptacle four scales
by nectary dish or nectary flowers flat
a (vs.
woody
amphisarcous
on and globose thick-walled fruits
indehiscent, ellipsoid
inserted in Quadrella)- to
it
oblong and few times ovoid, with seeds distinctively
with seeds capsular, dehiscent, linear or to
larger
(vs.
row
smaller, usually arranged in a single in Quadrella).
was known by two widely disjunct endemic species: Colicodendron yco from
previously
Colicodendron
and
Ecuador
dry western adjacent
and scabridum (Kunth) Seem., restricted to the forests of
eastern
Brazil, C.
American endemic both with whorled subsessile leaves
Peru. In this work, two additional South species,
added
and genus.
beautiful inflorescences are to this
&
bahianum nov
Colicodendron Cornejo H.H. (Figs. brazil. Amiga rodovia
X. sp. 1, 2, 7). Type: Bahia:
1. litis,
and
Cornejo
Revision of Colicodendron
litis,
Journal of the Botanical Research istitute of Texas 2(1)
78
13-40 3-7(-10) cm, coriaceous
narrowly x
with blades narrowly oblanceolate to elliptic, stiff,
subsessile,
sometimes
and shortly
and acute apiculate
chartaceous attenuate revolute the base, to
at
(dry),
(alive),
and
midvein, nerved glabrous
rostrate at the apex, shiny green (alive) with an impressed to often sulcate
midvein beneath,
prominent
and with glabrous or echinate-stellate
green strongly
above, paler a
(alive)
on
with 25 major upward-arching nerves each side of the
margin inconspicuosly thickened, 15 to lateral
wood,
and subterminal racemes, sometimes cauliflorous in older densely
terminal
midvein. Inflorescences
3-7
same
1-5 cm, bearing 25 flowers the time. Floral bracts linear,
axis short, stout, to at
stellate-echinate;
mm, 1.8-4 cm, the Sepals oblong-elliptic or
deciduous, densely echinate-stellate. Pedicels articulate to axis.
12-22 6-8 mm, mainly toward the base, ± acute apex, yellow
x abaxially keeled, at
lanceolate,
ellipsoid to
mm
4-6 by
hypanthium
brown deep, coated a subtet-
(when
rusty dry), stellate-echinate abaxially;
(alive),
2.5-4
x
ragonal 4-dentate nectary dish within, nectary appendages broadly deltoid to semiorbicular, 1.5
oblong
mm, bud, apparently inserted the edge of the nectary cup, to
glandular, Petals torsivae in at
stellate.
mm,
yellow
20-28 7-12 widely obtuse rounded apex, longitudinally nerved, sulfur
obovate-oblong, x to at
(when on both Stamens 30
rusty-brown sides. to
and erect anthesis (Mori, pers. com.), dull dry), stellate
at
expanded
on tomentose torus-
2-3 cm, geniculate in bud, glabrous, inserted a white-stellate-
filaments
42,
5-6
mm,
Gynophore 1.6-2 cm, glabrous; ovary oblong,
white
androgynophore, anthers 2
(alive).
like ca.
mm, with gynophores
x 2-3 longitudinally sulcate, glabrous; stigma sessile, capitate. Infructescences ca.
2-4
7-12 cm, subpendulous;
Amphisarcous x
2-2.5 x 0.3-0.5 cm, pedicels 3-3.5 cm. oblong-ellipsoid,
ca.
mm
mm
3-7 mature seeds 1.3-17 in diam.
wall thick,
fruit
Colicodendrum bahianum somewhat related to the equally yellow-flowered C. yco, is allopatric in the
is
more mesophytic bahianum can be distinguished
However, easily
dry Caatinga the north the C.
to (Fig.
7).
end branches 1A, sub-
by dense whorls arranged toward the of the terminal (Fig. vs.
of subsessile leaves,
The Colicodendrum
more inflorescences of
and spaced out along the branches).
opposite pedicellate leaves,
racemes and paniculate inflorescences, or rarely solitary axillary
bahianum are short dense larger longer,
(vs.
number
stamens
with sepals anthesis, higher of (30
flowers in yco\ bearing flowers erect reflexed) at
C. (vs.
mm
cm
mm), gynophores
4 glabrous (1.6-2
to 42 vs. 14 to 20 in C. yco), smaller anthers (ca. 2 vs. shorter
and pubescent)
3.5-5 cm, pubescent) glabrous ovaries.
vs. (vs.
more
Due bahianum seems morphologically closely related
whorled Colicodendron
the subsessile leaves,
to
&
Bunting
H.H. Ruiz differences see
G.S. (for
herein described Colicodendron valerabellum T.
to the also litis,
comment
under
the species).
latter
November and and August.
Phenology.— Collected in flower in February, August, January, fruit in
July,
hummingbirds
The by hermit (Webster 25010).
sulfur yellow flowers are visited
—
Unknown.
Cytology.
moist wet evergreen Atlantic
southern extension of the or
Distribution.—Eastern Brazil, restricted to the
mm)
2000 Bahia
southeastern
rain forests into State (Fig. 7).
(ca.
—
wood
human and
from both har-
under pressure agriculture
Habitat intense
Conservation concern.
is
comm,
Sep
bahianum Mori, 1979;
vesting, especially for so decorative species Colicodendrum (Scott per. "It
ornamental
Images potential species are
was one of the most beautiful species that I've ever seen"). of this
New Web
(NYBG,
Garden
York 2003).
Botanical
available the site
at
&
2580 UB, WIS).
BRAZIL. Bahia: Mun. Santa Cruz Cabralia, mata costeira, 15 Jul 1966, R.P. Belem R.S. Pinhtiro (F, Est.
Paratypes.
(CEPEC, DAV, Porto Seguro, Parq. Naca. Monte Pascoal, 15 Jan 1973 T.S. Santos 2692 (CEPEC, NY, WIS); ibidem, trail to
WIS)'. (fl.),
W
(CEPEC,
1971 Santos 1329
Jan T.S.
(fl),
NY
al 2283 (CEPEC, [NY photo in tt
hi et
(CEPEC, WIS).
Pinheiro2202 Enti
V-
73, R.S.
\A
fill
"M
C [CEPEC photo and fragm. WIS]).
at
Cornejo and
Revision of Colicodendron
litis,
Itabuno, prominent town CEPEC
for a in the Cacao region, where the herbarium mata
is located], Cascalheira, pluvial, 13
1
Colicodendron scabridum
2. (Kunth) Seem., Herald
Bot. Voy. 78. 1852.
(Fig.
7). Basionym: Capparisscc
B-W
NY!]; 10055-01 [B-W
at isolectotype: photo 9478
0! WIS!]).
at
&
&
DC,
Capparis angulata Ruiz ex
Pav. Prodr. 1:253. 1824. Colicodendron angulatum (Ruiz Hutch., Gen.
Pav.) 2:309.
Fl. PI. 15
PERU:
Type: Piura, Paita, Voyage de la Bonite, Jul 1836 M. Gaudichaud B
(fl, fr), s.n. (lectotype, designated here: 10 0242739!;
isolec-
B
totypes: 10 0242737! F F G
["misit 1841"]; 876547!, 894168!, F [fragm.] 609130!, [G photo 8472 WIS!],
at
P!).
Nomenclatural note.—The
herein
selected lectotype of Capparis housed
scabrida has with
at P, a label a correct
origin of the specimen, "in America equatoriale," and the number "19" added. The B
isolectotype Cap-
of
paris scabrida consist of a branch with a terminal inflorescence, with the two one and
leaves, inflorescence,
floral pieces detached and remounted. That
isolectotype, originally only was named
identified as Capparis,
Capparis Willdenow's nomen non
speciosa nudum,
(a Capparis =
speciosa Griseb.
Anisocapparis
[1879],
speciosa
&
[Griseb.] X. Cornejo H.H. which
litis), as reads the label is at the upper left corner of the photo 9478, now
(B-W
in a separate sheet 10055 -00 That has
label the following handwritten
information:
0). "Polyandria
subtus
glabris tomentosis
foliola elliptica. Habitat in America meridionali." The discussed
isolectotype
later
was
by
correctly
identified Eichler himself, in handwritten which
a label of determination,
reads: "Capparis
(Colicodedron) scabrida H.B.K."
m
cm
Multi-branched
shrubs and
to trees, to 10 25 dbh, low and widely
erect to spreading; stem
the
when gum
injured produces
a (Litde 6725); terminal branches complanate ±
or tetragonal, densely
stellate
throughout.
Leaves mm,
few
spirally-alternate, times 8-30
opposite-decussate,
petioles lacking
pulvini;
blade coriaceous when
(rigidly coriaceous (6-)8-23 2-7
dry), lanceolate oblong, x cm,
to acute rounded,
to
sometimes
emarginate,
apiculate apex, cuneate rounded, sometimes
at to retuse green
base,
at olive
(alive),
but drying
to a bright sulfur-yellow herbarium when
in
material, thinly stellate-tomentose young, soon
and smooth
glabrescent
or scabrous above, pale grayish or "ashen" greenish-yellow and
densely
pale
stellate-tomentose with prominent midvein
a beneath; (7-)12-23 on
nerves each
lateral side of the midvein.
Inflorescences simple, densely corymbose
racemes, and
terminal
solitary or lateral in the axils of leaves, or
branched, compounded
terminal racemes
complex,
into corymbose
stout, (sub)erect, panicles, 30 cm,
to
densely stellate, each raceme with only 1-4 open flowers end one 5-12
at its at time. Floral bracts filiform,
mm,
soon
deciduous. Pedicels 2.5-4 cm, densely brown and
stellate sparsely dendroid. Flowers dark brown
greenish-brown,
to densely Flowers asymmetrical,
stellate. in perfect flowers the androgynophore-gynophore
off-centered within the calyx cup, arching out and upward.
Sepal lobes broadly 10-15
ovate
to lanceolate, x
mm,
5-8
at anthesis cucullate-ascending, acute the apex, densely brown on both
at
stellate sides, eventually
caducous, broadly inserted on the rim of a wide bowl-shaped calyx cup, 9-13 x 6-8 mm, with
3-lobed
a
nectary cup hypanthium
coating
the within. narrowly
Petals broadly oblong 15-23
to ovate, or x
elliptic,
mm,
10-15
and
reflexed
outrolled at the tip at anthesis, widely cuneate to truncate at the sessile base, obtuse
rounded
to at the apex, white to cream or yellowish, sometimes greenish, on both Stamens
stellate sides.
mm
24 borne on
to 35, in a ring top of the short 2-3 androgynophore,
filaments (3.5-)5-7(-8) cm,
stellate
mm
and
dendroid, adnate each other 0.5-2 3-4 mm,
to for the base; anthers white Gynophore
at
(alive).
5-8
cm,
creamish-white, maturing
purple 4-6
to sparsely ovary ovoid 3-5
(alive), stellate; to ellipsoid, x
mm,
densely
stigma
stellate; truncate, Infructescences with gynophores 7-10
sessile. x 0.4-0.6 cm,
pedicels
2.5-3.5(-4) cm. Pseudoamphisarcum
oblongoid, ovoid, or 8-15 5-9
asymmetrical,
ellipsoid, slightly x
Research of Texas
Journal of the Botanical Institute 2(1)
80
when
mature exhibiting 8 longitudinal ± sulcate
umbonate
apex, densely
cm, pendulous, ± the stellate,
at
mm
6-10 pulp bright orange
wall thick,
4 major, indicating the valves); fruit fibrous,
(dried) lines (usually
cm,
1-2 x 1-1.5 sur-
subspherical-reniform, often strongly beveled, ca.
seeds 15
maturity, insipid; to 60,
at
many reddish-brown
by from the crustaceous testa;
rounded by orange hairs
densely sarcotesta infiltrated
a
embryo
yellow.
woody subwoody
with fruit-wall,
The amphisarcum indehiscent provided a thick, to
a simple, fruit,
is
maturity 1994:
upon removal and never in valves after (Spjut
which shape of sarcocarp, splits
the
retains
decompose and
on ground eventually
maturity the to
In Colicodendron scabridum, the fruits at fall
23, 37).
convex, unattached, persistent
thickly-subwoody, abaxially
the seeds, leaving usually four elliptical,
release
known
which any the types (Spjut
doesn't in of fruit
That type Colicodendron scabridum,
of fit
valves. fruit
which
amphisarcum) Cornejo (mod. nov),
Pseudoamphisarcum (pseudo + X.
proposed
herein as
1994),
is
campo Van den Eynden
ECUADOR:
names.— sapote de (Spanish, et
Sapote (Spanish, 6725),
Local Little
&
gomoso Steyermark
(Spanish, 54842)]
sapote
1999: sapote de perro (Spanish, Cornejo Bonifaz 7583),
42),
al.
Madsen
8521
64103).
Zapote de perro (Spanish, Acosta-Solis Mille 1940,
Brandbyge
Zapotillo (Spanish, 42789);
&
Zapote (Spanish,
Woytkowski
de perro (Spanish, 5672).
PERU: Diaz Baldeon sapote
Sapote (Spanish, 2374),
&
Simpson Schunke
(Spanish, 567).
Vargas Zapotillo
42),
The mature pulp Colicodendron scabridum strongly resembles the color
bright orange color of the fruit of
known [Bonpl]
{Quararibea cordata
"Sapote"
pulp Ecuador) well edible fruit called
of the of a locally (in
names
and Colicodendron
similar derived local of
Bombacaceae). This the origin the several
for
Vischer,
is
Thomas. This an endemic
one names, Pseudalopex sechurae is
"Sapote de of the local refers to
perro,"
known has distributional pat-
de monte" (Spanish) or "Sechura fox" (English), that a
canid as "Perro
locally
When
Ecuador and the food
scabridum in the dry coastal areas of Peru.
Colicodendron
tern similar to that of
(Bruning
scabridum
Colicodendron
the Pseudalopex sechurae facultatively eats the fruits of
really scarce,
is
Cornejo
2005).
1985;
Southwestern Ecuador,
Zapotillo (Spanish), a town located in a dry forest at the Province of Loja in
known
name name Colicodendron scabridum.
owes the local of the locally well
to
its
—
September December.
mostly
Flowering mostly between February October, fruiting to
to
Phenology.
m
A 4-6 60 per Within the nectary cup, three
produces 20 flowers anthesis night.
single tree high, to at
By
be produced 19:00 each droplet from below of each nectary lobe. 19:30 h,
nectar
droplets of start to at h.,
anymore,
morning have nectar
the nectary cup of nectar. But by the next at 6:00 h, the flowers don't
full
is
The during one night per flower
during nectar secreted
perhaps due the intense bat activity the night. is
to
Bahia de Caraquez,
The one night only and don't produce any scent (Cornejo field obs. in
flowers
only. live
Ecuador, July 2004).
ECUADOR. WIS, unpublished
n = Manabi: and 243 ms).
Cytology.— litis
8 [!Pazy, litis litis
element dry very dry or xerophytic
of to
Distribution.— Colicodendron scabridum a characteristic floristic
is
Ecuador
from
Ecuador and where dune former. This species ranges sea level in
western Peru, a
areas of is
m
and Peru south
2500 mountains Province in to
from Province Manabi, in the of the Loja,
the of to ca.
Ancash
Dept.
(Fig.
7).
by
scabridum are visited the
Colicodendron
and observations.—-The flowers of
Ecological interactions
field
when
common Dozens buzzing bees can be heard from under a tree
honey of
domestic bee, Apis mellijera L.
and around
The honey peaks in the flowers of
the nocturnal flowers are just about to open. bee's activity
The domestic bees re-visiting the flowers the
between 18:30 19:30 re-start
Colicodendron scabridum to h.
is
such
presumably pollen predator bees as Halictidae
morning, between 6:00 8:00 Nocturnal,
next as well to h.
and
While both bees touch the anthers the ovaries,
19:00 species of
the flowers h.
(Megaloptal), also visit at
many
From small bats frequent the
20:00 5:50
seems however that the true pollinators are bats. to h,
it
The
which exposed on long peduncles high above the foliage. bats
flowers of Colicodendron scabridum, are
from during than one second,
wings hovering while licking the nectar each flower less
beat their in the air
and
Cornejo Revision of Colicodendron
litis, 81
hummingbirds
The
as are apt to do. bats do not hold themselves -om the flowers as happens the African
to
Bignoniaceae, same
Kigelia ajricana [Lam.] Benth. (seen in Bahia de Caraquez, Ecuador, the where
locality
When
was by
Colicodendron scabridum studied the senior author). leaving the flower, the bat shakes the
inflo-
presumably
rescence, helping
pollination. interesting to note that only the individuals of Colicodendron
It is
growing
scabridum darkness by These
in total are visited bats. conspicuously have a higher productiv-
fruit
ity per tree than the remaining individuals located around the nocturnal lights along the roadsides in the
same
locality. Additionally, the flowers of Colicodendron scabridum are also visited by Amazilia
amazilia, a
who
hummingbird
new
visits the old (previous night's) flowers between 7:00 to 8:00 h, and the flowers just
open
about commonly
to at ca. 18:00 h. Finally, the flowers of Colicodendron scabridum are visited by wasps,
and
coleoptera
several species of ants (Cornejo field obs. in Bahia de Caraquez, Ecuador, July 2004).
—
The have been humans and
Uses. fruits reported as edible for as medicinal against colds and coughs.
commonly
They
are eaten by and donkeys Ceron Van den Eynden
cattle 22424, 1999:
(C. et et 42,
al. al.
&
wood
make
Cornejo while the used and Van
field obs.), is for fuel to handicrafts {Cornejo Bonijaz 7583,
den Eynden
et al. 1999: 42). Finally, a "glue" for paper can be obtained from the cut, gum-oozing ends of
the stems (Ceron
11876).
—
A
common
Conservation biology. characteristic species in the dry landscape, does not appear be
to
it
danger
in of extinction.
&
m,7Jan IK7
QCNE);
1994, Cornejo (GUAY,
X. Bonijaz ibidem,
C
i ;.
QCNE,
(MO,
11671 WIS);
al 22424 (QAP);JoahaciaMer
420 Sep
111,6 1991, C.Jossc 638
QCNE &
,ca.2m,22
Los
Playa Frailes Jan 1991, A. Gentry
[2]); C.
1
,
^!
9
^jr™^
L^7nmZ^Sl
1
Aug-8
Sep 1836 [voyage of H.M.S. Sulphur], G. W. Barclay 422 (BM
I
Pum
H. Eggers 14735 (GH, K, LE, M, US); ibidem, 1852, Andersson
12, Isla
s.n. (S);
50 m, 4 Nov J. Madsen 64103 (AAU); path Rio Hondo Cerro Yanzu
>9'S, ca. 1987 to
V
QCA,
(AAU[2], QCNE). El Oro: Parroquia San Antonio, Hcda. Montecarlo,
2:
2
Journal of the Botanical Research Institute of Texas 2(1)
W3
o o S,5m,7 Apr
Holm-Nielsen
WI! Huaq t,80 13 27 1980,
Anderssc (GB, Lorn:scrub fores , , L.
i i);
QCNE, W]
(MO, QAP,
1876
Loja:
I
L^^sTS
IB^!^^^X
eM^
Hl'S V,
V^T.'.
mT3°F QCA)
5^400 *eenLoja
(GB,
}] S
QCNE); Catama
above
Van den Eynden 615 (LOJA,
1996,
et a\.
:b
m
ndU NY, between mbo and La Toma, 1000-22
>i.o,^ cry dry slope, 1650 9 Oct .955, E. Asplu J077(B,K, LD, S); El Ta:
c
a.
]
,
G.HarUng 6065
(MO.QCNE.WIS; (S);Cata
>59,
Sep 199' etai:
6
a, Little
Apr
ma,ca. 1500 m, 24 19
3
(GH
54842
Steycrmflrfc
43,
J.
Gua
-651 (NY, WIS[2]); Rio
:?H
>S
L°r229^^^
E?
NWof
rB
il
m, 537
Knight 609 0* .400 R. Esplrwzfl CF
QCA, QCNE, V
6584 (NY,
Valdespino SoalZs^i
10 Jul
1
i,
X\^'
T^l
1
ckTwkhcaa
i^79<
'roads)
Quebrada Las Penas, betw. San Juan de la Virgen and
Lam-
>374 (MO).
La
Wright 3518 (MO);
ison
&J.
Maranon Celendin-Balsas 25 kn
Valley, rd.
&
Apr Baldedn 2784 (MO); Ce
Libcrtad: 15
Prov. Trujillo, Trujillo, 1 iz S.
A (MO, band
NY);
10965 (MO); ibidem, 100 m, 14 Jul 1973, Araujo 7978
:.
*s™^
7'maZ
GH,
tm&J. Wright 3352 G,
(F,
&
2-7).
Colicodendron valerabellum H.H. Ruiz Bunting, nov. (Figs.
G.S. sp.
3. T. i
litis,
damaged with one two unbranched stems from near the
Evergreen palmoid in plants or additional
treelets [or
and sometimes
flower-bearing
some decumbent and sending lateral-
base, of these forth 1 to 3, erect, leaf-
m
cm
3-5 dbh crowded young branches) dark-brown
and
long often (colonal?);
horizontal 5
shoots], to
grouped
cm,
in
echinate or echinate-stellate; stipules absent. Leaves subsessile to short-petiolate, to 1.5
nodes
crowded whorls the end of the terminal branches exposing mostly leafless internodes, of leaf inser-
at
4-9
cm,
3-6 narrowly (16-)20-50 x coriaceous
cm, blades oblanceolate stiffly
tions swollen, to elliptic,
ca.
and
Cornejo Revision of Colicodendron
litis,
and hard but somewhat
flexible chartaceous attenuate but rounded-retuse subcordate the
(alive), (dry), to
at
base, acuminate or acute to widely obtuse the apex, margin inconspicuosly thickened and
at revolute, lustrous
green (alive) with a impressed to sulcate midvein, prominently and coarsely rugose with the
bullate),
(i.e.,
blade surface between the deeply impressed midrib and main nerves prominently convex and smooth,
ap-
Journal of the Botanical Research Institute of Texas 2(1)
H.H. Ruiz & G.S. Bunting. A. Habit. B. A leaf blade. C. An inflorescence with three hermaphrodite flowers in
T.
litis,
i
An
male above them bloom and three beginning to wilt; and, above these, some 50 male flower buds. D.
postanthesis at base; four flowers in full
&
&
inflorescence bearing (sub)sessil flowerbuds. E-F. Immature fruit. (A. After photo of Bunting Trujillo 13120. B-D. litis etal. 30547, WIS. E-F. Ruiz
fane 4764, MY). A-E, ni; by T. Ruiz Z.
Villa F,