Revision of Spathacanthus (Acanthaceae)

14 Pages·1999·10.8 MB·

by  Thomas Daniel

Contr. Univ. Michigan Herb. 22: 33-46. 1999
SPATHACANTHUS (ACANTHACEAE)
OF
REVISION
Thomas
Daniel
F.
Department Botany
of
Academy
California of Sciences
Golden Gate Park
San 94118
Francisco, California
INTRODUCTION
Spathaccinthus BailL treated as comprising three species occurring in moist
is
and
wet Mexico and Central America. This genus of large shrubs
to forests of
among Acanthaceae by very large capsules
neotropical
small trees distinctive its
is
Few
among known
and seeds, both of which are the largest in the family. collec-
had been made There no previous
tions of the genus until relatively recently. is
two
comprehensive treatment of the genus; however, Daniel (1995a) treated of
Mexico.
the species in
The Spathacanthus have never been adequately addressed.
generic of
affinities
Lindau placed the genus in the tribe Asystasieae, which he characterized as
(1895)
having ''Rahmenpollen" 3-colporate and 6-pseudocolpate pollen with the
(i.e.,
pseudocolpi each mesocolpium fused near the poles to form pseudocolpal ellipses),
in
four stamens usually with dithecous anthers, and two seeds per capsule valve.
Bremekamp included Lindau's Asystasieae within subfamily Ruellioideae,
(1965)
Odontoneminae.
subtribe
tribe Justicieae,
&
Breedlove Smith
Examination of pollen in Spathacanthus {S. hahnlaniis:
& We
Hernandez Gonzalez
Hernandez. G, 1777. G. L. 1855,
Matuda
iflorus:
S, hoffi
&
Q
Marcial
Bremekamp's Odontonemi-
most
veals grains that are similar to others in (1965)
They and 6-pseudocolpate The two pseudocolpi
nae. are usually 3-colporate (Fig. 1).
each mesocolpium sometimes fused near one or both poles, but with equal
are
in
commonly
pseudocolpi
frequency they are not. Similar variation in fusion of is
encountered among American genera of Odontoneminae (Daniel 1998); however,
Unhke most American
two features of pollen in Spathacanthus are noteworthy.
Odontoneminae, the interapertural surfaces of the exine (excluding the pseudocolpi)
not conspicuously Rather, they vary from being foveolate to fossu-
are reticulate.
late to verrucate to rugulate. The absence of a conspicuously reticulate exine is
American Odon-
Spathacanthus other
derived character relative to
treated as a in
number was noted
toneminae. Secondly, variation in aperture in S. parviflorus
and
Pollen of Skutch 961 and Steyermark 46632 has four colpi eight
(Fig. Ig-i).
and
pseudocolpi; pollen of Matuda 1696 consists of grains with four colpi eight
and
pseudocolpi together with grains having three colpi and six pseudocolpi; pol-
&
Q
Marcial
M
number been encountered
Acanthaceae, variation of aperture within a species has
number
Even
aperture
only Stenandrium dulce (Daniel 1998). variation of
in
uncommon among American Odontoneminae; been noted
has
within genus
a it
is
33
CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME
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22
FIG. &
Pollen of Spathacandius. haluiianus (Hernandez Gonzalez
1. a. S. G, L. J855), apcrlural
&
view. haluiianus {Ilenuhidez Gonzalez
b. S. G. L. 1777), inlerapertural view. haluiianus
c. S.
(Wendt
et 2769). inlerapertural surface, hoffnuuuvi (Liesner
id. d. S. et 15444), inlerapertural view.
al.
e. hoffnicuuui {Skutch 3325). aperlural view. hoffmannii {Skutch polar
.S". S. 3325), v\c\w. parvijlonis
f. g. S.
{Matuda
1696), inlerapertural view. h. S. parviflorus {Matuda 1696), polar view of 3-aperturate grain,
i, .S". parviflorus (Sieyerfnurk 46632), polar view of 4-aperturate grain. Scale: a, bar = 10 b, d-h,
i, |.u-n;
bar = pm; - pm.
12.5 bar
c, 1.5
Odontonema
only in (Daniel 1995b). Additional sampling of pollen Spathacan-
in
thus clearly warranted.
is
The common
presence of four stamens not among Bremekamp's Odon-
is
The
toneminae. American
only other genus with four dithecous stamens
in this
subtribc ChalarotJiyrsus Lindau. Chalarothyrsus genus
is a unispecific occurring
is
in the dry forests of western Mexico. Pollen of Chalarothyrsus resembles
that of
.
SPATHACANTHUS
DANIEL:
35
1999
however,
Spathacanthus sometimes having pseudocolpal ellipses in the mesocolpia;
in
most American Odontoneminae by having
resembles that of other a reticulate
it
These two genera can be distinguished by the following couplet:
exine.
mm
m
homo-
Perennial herbs or small shrubs to 1.5 calyx small (to 5 long), 5-lobed, the lobes
tall;
mm mm
morphic; corolla red, 13-16 long; capsule 12-18 long; seeds permanently retained in
Chalarothyrsus.
capsule and partially fused to inner capsule wall.
mm
m
segments
Large shrubs or small trees to 8 calyx large (15-40 long), 2-parted, the 2
tall;
mm mm
entire or variously lobed at apex; corolla white to yellow, 23-95 long; capsule 40-85
long; seeds expelled from mature capsule, not fused to inner capsule wall. Spalhacanthus.
PHYLOGENY
A PAUP
was
(Swofford
Spathacanthus using 1993)
analysis of 3.1.1
cladistic
conducted reveal possible phylogenetic relationships of the species. Chalarothyrsus
to
was designated as the outgroup and ten characters were used (Table four of
1),
A
was
which were potentially informative. branch-and-bound search per-
(2-5)
ACCTRAN
Both
formed using equal weighting and unordered character states.
DELTRAN
and (delayed transformation) optimiza-
transformation)
(accelerated
A
which
methods were discussion of each character, includes infor-
tion utilized.
Odontoneminae known),
mation from other follows:
(if
among
Acan-
Calyx = green; = yellow. Colored calyces are rare
color: 1
1.
The
among Odontoneminae yellow
and bright
thaceae general in particular.
in
when mature unusual and
calyx exhibited by parviflorus the corolla highly
S. is is
probably the relatively small flowers of this species.
attracts floral visitors to
=
Calyx = spathaceous and with major divisions equal; spatha-
division: 1
2.
- and
and major unequal; 2 not spathaceous equally 5-parted.
ceous with divisions
name
proximally
The from which the genus takes a calyx that fused
feature its is is
unique
two segments This type of calyx
and comprises large, spathelike distally. is
Spathacanthus among Odontoneminae. Whether the divisions of the major
to
The pentamerous and deeply
segments are equal or unequal varies in the genus.
divided calyx of Chalarothyrsus typical of other Odontoneminae. This multi-
is
character could be treated as two binary characters: calyx lobes fused into
state
two major segments, and major
two major segments calyx lobes not fused into
vs.
The
calyx segments equally divided vs. major calyx segments unequally divided.
character would apply solely to the ingroup.
latter
mm
mm
=
Corolla length: = short (15-29 long); 1 long (46-97 long).
3.
among
both hahnianus and hoffmannii are exceptionally large
Corollas of
S. S.
Odontoneminae.
known
Corolla = white; ^ yellow; 2 = red. These and other colors are
color: 1
4.
and
among Odontoneminae. Because character likely very labile
for corollas this is
among
outgroup not represented species of Spathacan-
because corolla color of the
is
not possible to determine directionality of the states represented fl/?non.
thus,
it is
mm
mm
=
= long (8-20
Length of lobes of lower lip: short (1-6 long); 1
5.
How-
long). possible that this character linked to corolla length (character 3).
It is is
have
measurement ranges noted herein, there are taxa that short
ever, using the
mcvaughu and
with long lobes Mexacanthus T. F. Daniel) long corollas
corollas
(e.g.,
Odontoneminae.
anomala Leonard) elsewhere
the
with short lobes Dicliptera in
(e.g.,
Number = = 3^. Triaperturate pollen
of apertures in pollen: 3; 1 is
6.
among Odontoneminae. See above concerning pollen of
widespread discussion
Spathacanthus
CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME
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123456789
Table Data
matrix for species of Spallmcanfhiis and the outgroup Chalarothyrsiis.
1.
011110011
001010011
10
110001111
hahn'uuuis
S.
0202
0000
hojjinannii
S.
parviflorus
S.
Chalarothyrsiis
1
Polar diameter of pollen: = 54-67 = 39-51
7. jim; \im. In Chalarothyrsiis
1
the polar diameter of pollen varies from 54-59 |im and overlaps measurements
of
this diameter in hahniamis. Although large pollen might be correlated with
S.
large corollas within Spathacauthiis, this obviously not the case the outgroup
is in
where
the corollas are relatively small and the pollen
relatively large.
is
Exine = =
8. surface: reticulate; 1 foveolate to verrucate to rugulate. Exine
of interapertural surfaces in Chalarothyrsiis and most other Odontoneminae
is
conspicuously
reticulate. In species of Spathacanthus, the exine varies from foveo-
late to fossuiate to verrucate to rugulate.
mm
mm
9. Capsule length: = short (4-30 long); = long (40-90
long).
1
Capsules of Spathacanthus known among American
are the longest Odontonemi-
nae (which otherwise vary from mm).
in length 4 to 27 Capsules of Chalarothyrsiis
mm
vary from 12-18
in length.
Seed = =
10. fusion: seeds not fused to capsule valves; seeds fused capsule
1 to
Fusion
valves. of the seed coat to the inner wall of the capsule, observed
as
in
Chalarothyrsiis, has been reported previously
in the family only the unispecific
in
Mexican genus Aphanosperma
(Daniel This genus
1988). latter also treated
in
is
Odontoneminae,
the but has pseudopapilionaceous corollas and an androecium
it
two
of stamens. Therefore, does not appear be
to particularly closely related
it to
Chalarothyrsiis.
The
phylogenetic analysis resulted most parsimonious
in a single tree (Fig.
2)
RC
HI No
of 12 steps (CI - = RI = =
1.0, 0, 1.0, 1.0). differences in character state
ACCTRAN
DELTRAN
changes from
resulted the optimization. Even one
vs.
if
of the possibly linked characters noted above and from
3 deleted the
(i.e., 5)
is
same
analysis, a tree with the topology (but with one less step) obtained. Divid-
is
ing character two
2 into binary characters, as noted above, also results in a single
same
tree with the topology.
On
the basis of character states in other Odontoneminae. most characters
at
^
the base of the tree in Fig. 2 can be unambiguously mapped onto one
of the
Given
clades. absence elsewhere
their in the subtribe, the exceptionally long cap-
sules the two-parted and spathelike calyces and the non-reticulate exine
(9),
(2),
of pollen grains likely are synapomorphic ^or Spathacanthus. Permanent
(8) reten-
tion of seeds in the capsular valves (10) would appear to represent a derived
As
condition in Chalarothyrsus. noted above, corolla color cannot be polarized on
the basis of character states occurring other Odontoneminae. The change
in
to
red corollas Chalarothyrsus mapped)
in (as could with equal hkelihood have been
shown
change
as a to white corollas Spathacanthus.
in
Within
Spathacanthus, hahnianus and
S. S. hoffi
-77,
has white
corollas like hoffmannil but sp
S.
ments
hahnianus,
like
S. sister to this clade.
is
—
SPATHACANTHUS
DANIEL:
1999 37
4(1)
S.hahnianus
3(1) 5(1)
2(0)
hoffmannii
S.
1(1) 6(1) 7(1)
S. parviflorus
4(2) 10(1)
Chalarothyrsus
FIG. Cla(Jogram of hypothesized phylogenetic relationships of species of Spathacanthiis\ Cha-
2.
larothyrsus the outgroup.
is
TAXONOMY
Spathacanthus 1891. Type: Spathacanthus hahnianus
BailL, Hist. 10: 444. Baill.
pi.
Shrubs or small trees with cystoliths, stems sometimes with conspicuous pro-
Leaves
truding or blisterlike lenticels. opposite, subsessile to petiolate, entire.
racemes mostly
Inflorescence of terminal dichasiate (or thyrses); dichasia opposite,
pedunculate), subtended by Bracts opposite, green,
1-flowered, sessile (to a bract.
+ homomorphic
inconspicuous, Flowers subtended by 2 bracteoles, pedi-
entire.
pedicels usually enlarged Calyx green or yellowish, spathaceous,
cellate, in fruit.
beaked in bud, equally or unequally divided into 2 segments, segment comprising
1
2 fused lobes, apically entire to 2-lobed, the other segment comprising 3 fused lobes,
apically entire to 3-lobcd. Corolla white or yellow, externally glabrous (although
margins of lobes sometimes ciliolate), tube expanded distally into a ± distinct
throat, limb bilabiate (or appearing ± regular), upper lip deeply 2-lobed, lower lip
Stamens
deeply 3-lobed, corolla lobes imbricate (ascending cochlear) in bud.
4,
didynamous, inserted at base of corolla throat, included in corolla tube, anthers 2-
appendages,
thecous, thecae equal in size, parallel, equally inserted, lacking basal
direction of dehiscence (as observed in 2 specimens): anterior pair dehiscing toward
upper hp and posterior pair dehiscing toward lower pollen prolate spheroidal
lip;
to euprolate, 3 (-4)-colporate, 6 (-8)-pseudocolpate, pairs of pseudocolpi some-
times fused near or both poles thereby forming pseudocolpal arcs or ellipses in
1
staminodes
mesocolpia, exine foveolate to fossulate to verrucate to rugulate;
0.
Capsule
Style included in corolla tube, glabrous, stigma 2-lobed, lobes equal.
very glabrous. Seeds homomorphic, borne on retinacula,
stipitate, large, flat-
4,
tened, lacking trichomes.
Spathacanthus a genus of three species occurring in moist to wet forests in
is
southern Mexico and Central America.
— — —
CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME
38 22
KfY
TO THE Species of Spathacanthus
mm
1. Calyx al anlhesis equally or subequally divided into 2 prominent segments; corolla 73-97
mm
50-89
long; capsule long; Costa Rica.
S. hoffniafinii.
^. Calyx at anthesis unequally divided into 2 (rarely more) prominent segments; corolla 23-70
mm mm
40-62
long; capsule long.
mm mm mm
2. Corolla white, 23-29 long, throat 5-7.5 in diameter, limb 5-8 in diameter
mm
with lobes 1-3 long; calyx yellow prior to fruiting; Mexico and Guatemala on the
Pacific slope. parviflonis.
S.
mm mm mm
Corolla yellow, 46-70 long, throat 9.5-16 diameter, limb 22-45 diameter
2. in in
mm
with lobes (5-) 8-15 long; calyx green prior to fruiting; Mexico, Guatemala, and
Honduras on Gulf and Caribbean
the slopes. hahniamis.
S.
Spathacanthus hahnianus Jan-Feb
Baill., Hist. pi. 10: 444. 1891. Type: Mexic^o.
Veracruz: Misanlla, forct de la montays Santa Rita, 3 Jul 1866, L. Hahn
349
(holotype: P!); see discussion in Daniel (1995a: 130).
Macfadyena Donn.
simpUcifolia Sm., Bot. Gaz. (Crawfordsville)
16: 198. Jul
&
1891. Spathacanthus simpUcifolius (Donn. Sm.) Lindau ex Bureau K.
Schum.
in C. Martius, Fl. bras. 8(2): 294. 1897 [combination also proposed
by Leonard, Proc. Biol. Soc. Wash. 50: 16. 1937]. Spathacanthus donncll-
Lindau Donn.
snihhii ex Sm., Bot. Gaz. (Crawfordsville) 20: 293. Jul 1895,
nomen name
(new M. synonym
superfl. for smiplicifoUa but latter cited as
name
and dropped
''specific as inappropriate"). Spathacanthus donnell-
Aug
nomcn
smithianus Lindau, Herb.
Bull. Boissier 371. 1895, superfl.
3:
{M. simpUcifoha cited synonymy). Type: Guatemala. Alta Verapaz:
in
Aug
Pansamala, 3800 1886, //.\'o/7 Tiirckhcun 1030 (holotype: US!;
ft,
GH!
M!
K!
isotypes: US!).
&
Moc,
Rucllia macrocarpus Scsse mexic, non
Fl. ed. 2. 148. 1894, Ruellia
niacrocarpa Wall.. 1830. Type: not designated; see discussion Daniel
in
(1995a:
130).
m
Branched many Young
shrubs or small trees to 8 often with roots.
tall, stilt
somewhat
stems quadrate compressed, pubescent nodes where
to (especially
at
trichomes often form dense tufts) with flexuose to antrorsely appressed eglandu-
mm
mm
trichomes soon Leaves
lar to 1 long, glabrate. petiolate, petioles to 45
mm
long, blades broadly 45-250
elliptic to elliptic to obovate-elliptic, (36-) long,
mm
17-143
(9-) wide, 1.3-3.1 (-4) times longer than wide, (rounded to acute to)
acuminate
at apex, acute at base, adaxial surface glabrous, abaxial surface gla-
brous or pubescent along major veins with flexuose appressed eglandular
to
tri-
mm
chomes
to 0.5 long, margin glabrous. Inflorescence of terminal racemes or a
cm
basally branched panicle of racemes to 5 long (excluding flowers), rachis nearly
glabrous or pubescent with flexuose-appressed eglandular trichomes, often enlarged
in fruit; dichasia -flowered, sessile (sometimes becoming subsessile in fruit with
1
mm mm
mm
peduncles
to 2 long). Bracts triangular to subulate, 1.5-6 long, 1-2.3
wide, abaxial surface nearly glabrous or pubescent Bracteoles
like rachis. triangu-
mm mm
subulate 1-6
lar to to linear-lanceolate, long, 0.8-1.5 wide, abaxial sur-
face nearly glabrous or pubescent Flowers
like rachis. subsessile to pedicellate,
mm mm
pedicels to 7 long, 3-3.7 diameter glabrous. Calyx green,
in in fruit, (18-)
mm mm
22-43 8-14 diameter (measured beaked
long, in abaxially glabrous,
flat),
mm
bud beak
in with to 2 long, divided into 2 elliptic to ovate-elliptic segments
mm
mm
with unequal incisions 16-30 long on 1 side and (0.5-) 3- 15 long on the
—
SPATHACANTHUS
DANIEL:
1999 39
(when
other, longer incision 0.56-0.87 times as long as calyx shorter incision less
mm
than long the calyx then appearing particularly spathelike), segment gen-
1
1
mm
segment
erally with 2 triangular lobes 0.4-2 long, other generally with 3
mm
triangular lobes 0.4-4 long, lobes rarely longer and calyx then appearing
unequally 5-lobed. Corolla yellow (white fide Ventura 19361 from Veracruz and
mm
Holmes 4392 from Honduras), 46-70 long, narrow proximal portion of tube
mm mm
13-20 long, 2.5-4 in diameter, densely pubescent internally, throat 19-33
mm mm mm
9.5-16 diameter widest expanse, limb 22-45 diameter,
long, in at in
mm mm mm
upper 12-19 long, lobes 9-10.5 long, 4.5-8.5 wide,
lip elliptic, (5-)
mm mm
lower lip 15-18 long, lobes linear-elliptic to elliptic, 8-15 long, (3.5-) 7-8.5
mm mm
(-11) wide. Stamens with longer pair 16-18.5 long and shorter pair 12-
mm mm
14.5 long, thecae 3.3-4.2 long; pollen subprolate to euprolate, 3-colpo-
rate, polar diameter 56-64 )jm, equatorial diameter 42-43 [im, P:E = 1.31-1.51,
mm
exine foveolate to fossulate to rugulate. Style 28-35 long, stigma lobes 0.7
mm mm mm mm
Capsule 43-62 27-34 head 25-28
long. long, stipe long, long,
mm
retinacula 7-10 long. Seeds subcircular to somewhat squarish to deltate in
mm mm mm
low
outline, 6,3-9 long, 5.7-8 wide, 2-2.3 thick, surfaces with verru-
bumps.
cose ridges or
Illustrations. Fig. Bot. Gaz. (Crawfordsville) 16: pL 18, 1891.
3;
December-June.
Phenology. Flowering: throughout the year;
fruiting:
Distribution Mexico (Chiapas, Oaxaca, Puebla, and Veracruz), Gua-
(Fig.
4).
temala (Alta Verapaz), and Honduras (Yoro); floodplains and along arroyos
in in
lowland lower montane mesophytic montane and
rain forests, rain forests, forests,
oak 100-1350 m.
forests;
This species of the Gulf slope of southern Mexico and the Caribbean slope of
northern Central America was not included by Standley account of
(1926) in his
known
from few
the ligneous flora of Mexico. Until recently, the species w^as
A
known
Mexican and poorly Central America. nearly
collections, in
it is still
E
specimen from Guatemala (mountains on road Tamahu,
sterile of Tactic to P.
Standley 71343 at F) was annotated as this species by Leonard and cited by Gib-
The
son condition of the specimen does not allow for identification
(1974). its
my am
Mac-
using key, and therefore hesitant to include Thus, the type of
I it.
known Gua-
fadyena simplicifolia represents the only occurrence of this species in
A
known
temala. single collection, noted as having white corollas, also from
is
Honduras. White were noted on Ventura A. 19361 from Veracruz.
corollas also
It
remains be verified whether notations on these two specimens accurately por-
to
on some
tray a difference in corolla color individuals of the species.
The racemose inflorescences of hahnianus sometimes become more or less
S.
by between and
thyrsoid elongation of the region the bracts bracteoles
in fruit slight
forming a peduncle).
(i.e.,
km
Addfuonal Specimens Examined. Honduras. Yoro: 16 from Yaruclia [=Yaruca?] on
ca.
Quebrada de Oro lo Cerro Biifalo, W. Holmes 4392 (NY, TEX). Mexico. Chiapas: Mpio. Berri-
N &
km
ozabal, 13 of Berriozabal near Pozo Turipache and Finca El Suspiro, Breedlove A. Smith
Z>.
DUKE, MO, NY, RSA, TEX, & Thome ENCB,
21618 (DS, US, WIS), D, Breedlove 30868 (DS,
F, R.
MO); Mpio. Berriozabal, a 12 km N de Berriozabal, E. Martinez * M. Soto M-2424] (CAS,
S.
MEXU);
Mpio. Berriozabal, La Aduana, cerca de Rancho Flor de Corazon, hacia Cairo, E. PaJacios E.
el
MEXU).
1726 (CAS, CHIP, Oaxaca: Mpio. San Felipe Usila, Campamento Cerro Verde, carr. para
(MEXU);
Arroyo Tambor, Calzcida al 16596 Mpio. Santiago Lachiguiri, Dislr. Tehuantepec, Cerro
/. et
km SO &
Campos
de Buenavista, 2 de Crucero Buenavista, 16°45'N, 95°30"W, A. R. Torres 4103
V.
km E
(MEXU); Uxpanapa Region, between Esmeralda (17 of Sarabia) and Rfo Verde, 1.1 mi S of
CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME
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22
, I
L
^
\
1
L i_ }
'
1
^ r
1 V I
1 } k '
4
1
;
e
in^
(11 fK\
SI
'iiii
k
FIG.
SpatliaccuUhus luih/iianus 33 from Flora of Chiapas, used with permission),
3. (fig. Pi. 4^ a.
Hahil. h. Infloreseenee node. c. Flower, d. Calyx spill open. e. Corolla split open showing stamens,
f.
mm; mm; mm;
Capsule, g. Seed. Scale: a, bar = 2 cm; b, bar = 3 c, bar = cm; d, bar = 8 e, bar = 6
f,
I
mm. & &
bar = cm; bar ^ (Based Hcrndudez Gonzalez Hernandez
2.5 on; a, b, G. L. 1855] c-e, G.
I g,
Gonzalez Wendt Drawn
L. I777\ 3773.) by Ellen del Vallc.
f, g, ci al.
Esmeralda, IT^HTN, 94M5'W, Croat D. Jhmnon 63303 (BM, CAS, ENCB, MO, TEX); Mpio.
7\ c^
N
Sta. Marfa Chimalapa, Arroyo Matzpac, de Sta. Maria por vereda Rfo Verde, 16°58'N,
la al
& MEXU, MO,
94^41 'W, I/erndndez G. Gonzalez 1777 (CAS, TEX); Mpio. Maria
//. C. L. Sta.
Chimalapa, Rio Verde por la vereda a la cabecera, ca. 7 km N de Sta. Maria, 16"59'N, 94''40'W, //.
Hernandez G. & C. Gonzalez L. IS55 (CAS, MEXU, MO, TEX); Mpio. San Felipe Usila, Nueva
—
—
SPATHACANTHUS
DANIEL:
1999
41
FIG. Distribution of hohniamis (dots) and parviflorus (open squares).
4. S. S.
Santa Flora, R. de Santiago & A. Maria 247 (MEXU); Mpio. Guevea de Humboldt, Dislr. de
II.
N
km
Tehuantcpec, recorrido La Cumbre-arroyo seco, 13.4 de Guevea de Humboldt, 16°52'N,
—
C
&
Campos (MEXU).
95°25'W, R. Torres A, 13897 Puebla\: Tzitzinapan, Yancuictlalpan, Cuetza-
V.
&
(MEXU); (MEXU);
lan, F. Basurto R. Patron 454 cerca de Atexcaco, Gold 324 Patla/Necaxatal,
Z).
C
Troll 325 (M). Veracruz: Colonia Melchor Ocampo, Cerro del Chiquihuite, C. Conzatti 4 (GH);
Km
Mpio. Hidalgotitlan, 4 camino Cedillo-La Escuadra, IT^OO'N, 94^40'W, Dorantcs al 3766
/. et
Km
(ENCB);
Mpio. Hidalgotitlan, 4 del camino Hnos. Cedillo-La Hulera, Dorantes 4058
/. et al.
SW
(ENCB); Nuevo 94°2rW, &
de Esfuerzo en cascada, Minatitlan, H'^IO'N, Rivera H.
la
/. F.
Gomez km E
8 (CAS); 3 de Rfo Chalchijapa por carretera Sarabia-Cedillo, Jesus Carranza, M.
la
km O
Vazquez et al 1584 (ENCB); Mpio. Hidalgotitlan, Rio Soloxuchil, 1.5 del Campamento Hnos.
MEXU);
(ENCB,
Cedillo, 17°16'N, 94^36'W, M. Vazquez 1611 Mpio, Atzalan, La Calavcra,
et al. F.
MEXU);
(ENCB,
Ventura A. 11601 Mpio. San Andres Tlalnchuayocan, Ventura A. 19361 (ENCB);
F.
Mpio. Minatitlan, Rio Uxpanapa, cerca del limite con Oaxaca, 17°irN, 94°08'W, Wendt 2769
T. et at.
MEXU, MO,
(CAS, TEX); Mpio. Hidalgotitlan, Arroyo Mazate S de Ejido Agustin Mclgar SE
al al
MEXU,
MO,
de Paso de Moral, 17n3'N, 94°35'W, Wendt al 3773 (CAS, TEX); Mpio. Minati-
T. et
O
km &
tlan, 4.5 de Uxpanapa, sobre terraceria a La Laguna, 17''12'N, 94°14'W, Wendt Almaraz
T.
I.
G. 4194 (CAS, MEXU, MO, NY, TEX).—State Undetermined: without locality, M. Sesse & Mocirio
J.
(MA, MA,
302 US); M. Sesse 2146 US).
et (F,
al.
Spathacanthus hoffmannii Lindau, Bull. Herb. Boissier 370. 1895. Type: Costa
3:
C
Aug
RiCA. Alajuela: prope Aguacate, 1857, Hoffmann 842 (holotype:
GH!
B, destroyed, photos: F! US!).
m
Shrubs up
to small trees to 8 trunk (often with multiple trunks) to 10
tall,
cm
Young
somewhat
in diameter. stems quadrate to flattened, glabrous (or the
nodes pubescent with brownish, flexuose eglandular trichomes). Leaves subsessile
mm
to petiolate, petioles to 15 long, blades eUiptic to elliptic-obovate to obovate,
—
HERBARIUM VOLUME
CONTR. UNIVERSITY OF MICHIGAN
22
42
mm mm
89-285 long, 25-109 wide, 1,8-5.2 times longer than wide, abruptly acumi-
and margin
nate to apiculate at apex, acute to attenuate at base, surfaces gla-
mm
raceme
brous. Inflorescence a terminal to 35 long (excluding flow^ers), rachis
young
pubescent like stems; dichasia 1-flowered, sessile. Bracts triangular to lan-
mm mm
2-8
ceolate, long, 1-1.3 wide, abaxial surface glabrous. Bracteoles lance-
mm mm
2-5 1-L5 Flowers
subulate, long, wide, abaxial surface glabrous. pedicel-
mm mm
25-
late, pedicels to 9 long, 3.1 in diameter in fruit, glabrous. Calyx green,
mm
mm
40 long (somewhat accrescent 10-16 diameter (measured
in fruit), (8-) in
mm
beaked bud beak
abaxially glabrous, with 0.6-1.2 long, equally or
in a
flat),
subequally divided into 2 ovate to elliptic to obovate-elliptic segments, incisions
mm
when
11-22 (-28) long (rupturing further capsules dehisce), 0.34-0,73 times
as long as calyx, segment apically entire, other segment apically entire or 2-
i
mm mm
73-97
lobed, lobes triangular, to 1.5 (-5) long. Corolla white, long, nar-
mm mm
row proximal portion of tube 18-30 long, 2-5 internally pubescent,
in,
mm mm mm
35-50 9-18 diameter near midpoint, limb 30-42
throat long, in in
mm mm
diameter, upper lip 15-26 long, lobes ovate to elliptic, 10-20 long, 11-18
mm mm mm
wide, lower 18-25 long, lobes ovate to 15-20 long, 12-16
lip elliptic,
mm mm mm
wide. Stamens with longer pair 24-34 long and shorter pair 18-28 long,
mm
thecae 4-6 long; pollen subprolate to euprolate, 3-colporate, polar diameter
65-71 jam, equatorial diameter 44-56 |am, P:E = 1.16-1.62, exine fovcolatc to
mm mm
44-59
foveolate-verrucate to subrugulate. Style long, stigma lobes 0.3-0.6
mm mm mm
Capsule 50-89 20-40 head 27-39
long. long, stipe long, long, retinac-
mm
7-12 Seeds somewhat shaped subcordate
ula long. irregularly to to subelliptic
mm mm mm
in outline, 7.5-13 long, 7.5-11 wide, 3-5 thick, surface with shallow
subconcentric ridges (especially toward margin).
Illustration. Fieldiana, Bot. (n.s.) 18: 19, fig. 17, 1986.
Phenology. Flowering: January-March and May-October; Decem-
fruiting:
ber-March, and May-September.
Distribution Costa Rica (Alajuela, Cartago, Guanacaste, Heredia,
(Fig.
5).
Limon, Puntarenas, and San Jose); along streams in regions of humid forests, rain
forests, and cloud forests on both the Caribbean and Pacific escarpments; 80-1700 m.
more
Based on the cladistic analysis Spathacanthus hoffmannii closely related
is
The
to hahnianiis than to parviflonis. former two species both have long
S,
.S*.
corollas with the lower lip terminating in relatively long lobes. Like parviflonis,
S,
however, hoffmannii has white
corollas.
S.
Additional Sprcimens Examined. Costa Rica. Ai.ajuela: Esl. Biol. Rcserva Foreslal de San
Ramon,
valley oC Rio Lorencilo, 10^13'N, 84"37'W, W, Burger al 12439 Bajos dc Jamaica],
(F);
et
Reserva de San Ramon, Chacon 1768 (CR, DUKE); Buena Visla, San Carlos, D. Cox 5 (DUKE);
/.
NW
Finca Los Ensayos ca. 11 mi of Zarccro, T. Croat 43516 (MO); San Jose de Naranjo, Garcia 1
/.
E
Ramon,
(CR, Rfo San Lorencito, de Reserva Forestal de San Gdinez-Laurito 10263
F); Ifniite la
./.
(CR, F); Est. Biol. Res. Foreslal de San Ramon, valley of Rfo Lorencito on Caribbean slope, 10°13'N,
&
84°37'W, Gornez-Laurlto K. Swangcl 12439 (CR); Rcserva Forestal de San Ramon, Rfo San
./.
MEXU,
Lorencito, 10°14'13"N, 84°36'22"W, G. Ilerrera Ch. 362 (CAS, US); Buena Vista de San
Carlos, L, HoUlridge 6790 (CR); Quebrada Lajas, Finca Los Ensayos, Buena Vista de San Carlos, A.
MO, Lem
Jimenez M. 2319 (BM, CR, NY); near road Laguna Hule, 10°17'N, 84n2'W, 3243
F, to R.
MO); & (DUKE);
(CR, along Rfo Sarapiqui, mi S of Cariblanco, R. Primack D. Stone 44
F, 1
&
Cordillera de Tilaran, Rfo La Balsa, 10^15'N, 84'^3rW, V. Ramirez Q. Jimenez 273 (NY); rd to
(DUKE,
Colonia Virgen del Socorro, barranca of Rfo Sarapiquf, 10°15'N, 84^10'W, W. Stevens 13547
F, MEXU). Cartago: above La Suiza, ca. 0.5 mi behind town, S of Turrialba, G. De Wolf 307 (K,
NY); carrctera entre Turrialba Siquirres, a vera del Rfo Chitaria, Gomez-Laurito 6801 (CR,
y la
J.
USJ); along Rfo Chitarfa, R. Lent 1414 (F, US); Cerros de La Carpintcra, 9^53'N, 83"59'W, R, Lent