Table Of ContentON
HAPLOPHYTON
NOTES
MISCELLANEOUS
(APOCYNACEAE: PLUMERIEAE:
HAPLOPHYTINAE)
KIRK WILLIAMS
JUSTIN
Department Botany
of
Texas
University
of
TX
78713. U.S.A.
Austin,
ABSTRACT
by various
/pic
;
examination of herbarium specimens coupled with fieW observations Chi<
3se ir1
shoul
Hc,plol^hyto„
,at
tribe,
aplophytinae. Distribution maps and a key to the species are provided.
Words: Haplophytmae, Alstominae Apocynaceae
Hapbphyton, Alstoma,
EY
,
De CandoUe (1844) described Haplophyton as a monotypic genus based
upon H. cimictdum. Benson (1942) added a variety to the species, var. crooksii,
when
later elevating to the rank of species (Benson 1943) material of
it
mature seeds was examined. Benson (1954) regarded H. crooksii as a dis-
taxon because of smaller leaves and smaller discontinuously ridged
tinct
its
Leeuwenberg
Various authors gohnston 1990; Kartesz 1994; 1994;
seeds.
and recognized only
Pichon however, disregarded these differences
1950),
TEX
My
exammation specimens housed LL, coupled
H. dnnctdum. of the at
new
with observations in Chiapas, Mexico, however, has uncovered
field
two
evidence that strengthens the recognition of distinct species.
The subfamily Plumerioideae because
genus Haplophyton placed in the
is
of undifferentiated free anthers and mdole alkaloides. These characters,
its
mono-
particular the indole alkaloides, are regarded as indicative of a
in
phyletic lineage (Leeuwenberg 1994). In addition, sinistrorse aestivation
bud
of the corolla (overlapping of the petals in to the left) distinguishes the
Plumerioideae from the Apocynoideae, which
has dextrorse aestivation
(over-
lapping bud
of the petals in to the right) (Leeuwenberg 1994). The
aestivation oiHaplophyton, however, has been under considerable question.
De
Candolle
(1844) described Haplophyton
as sinistrorsely contorted, while
Bentham
and Hooker
(1873) listed the taxon having dextrorse
as a
Pichon
aestivation. (1950) also pointed out that Haplophytou had
a "curi-
Woodson
ous" dextrorse aestivation. (1938) on the other hand reported
a
sinistrorse aestivation Haplophyton, did Standley and Williams
for as
(1969),
and and
Correll Johnston Benson
(1970). (1942, 1943, 1954) did not de-
An
scribe the aestivation. examination of herbarium specimens, however,
reveals that the above reports are correct. The southwestern United
all States
and northern Mexican members
(Fig. o{ Haplophyton have an
aestivation
1 )
to the while the Guatemalan and southern Mexican members
left,
(Fig. 2)
have an aestivation to the right. The correlation of this character with the
allopatric distribution of the two populations provides strong evidence
that
My
Haplophyton comprised two
of
distinct
is species. field studies in El
&
Chorreadero, Chiapas
{Williams Plum, 95-34) confirm that southern popu-
When
Cando
de 1844) d^escribed Haplophyto? ikidum he reported
a
"
type a spec: n from Tehuantepec,
•
'
Oaxaca. Topotypes from
this
regi(
Because have not had an opportunity to examine the type, aestivation
its
I
remains present undetermined. suspect, hov^^ever, that to the right
at I it is
from specimens
and that the reports of a sinistrorse aestivation are either
examined from the north or are the result of the presupposition that all
Plumerioideae flowers have aestivation to the
left.
now
Differences in the corolla tube and calyx dimensions have not until
my
between two but
been discussed segregating factors the species, ex-
as
amination of herbarium specimens shows that H. cimicidum possesses a longer
An
examination
tube than H. well smaller calyx lobes.
corolla crooksii, as as
and supports
of the seeds, in particular their size surface structures, also
coma which
that seems to tentatively hold true the color of the is typi-
is
golden yellow H. cimiadim and charcoal grey in U.
cally in crooksii.
The populations Guaymas, southwestern Sonora, superficially resemble
in
them
H. cimicidum in leaf size; accordingly Benson (1943, 1954) recognized
A
however, shows
such. close inspection of these individuals, that their
as
mm
6-8 them
aestivation to the left and that the seeds are long, placing
is
with H.
crooksii.
With
sinistrorse aestivation, larger calyx lobes, smaller corolla tube,
its
and geographical range, seems evident that
smaller seeds, isolated
it
SiDA 1995
16(3)
Haplophyton constitutes distinct element from H. cimkidiim and,
crooksii a
two
consequently, necessitates the recognition of species for Haplophyton.
mm mm mm
8-10
3-5
wide; corolla tube long; sepals long,
mm
8-1
the length of corolla tube; seeds long, vertical
ri
1
m
bud overlappmg 1-35
Petals of the to the leaves (50)
left;
1
.
mm mm mm
6-8 4-1
wide; corolla tube long; sepals long,
1
I
mm
6-8
equal to half the length of corolla tube; seeds long,
\
western United and northern Mexico
States
(TEX)
El
(
iTL\)
34
i
km
(TEX)
^2
S of
/
(TLX) 9-
en
/ 01 irc is
(TLX)
J59 Knt I^l
D D
Haplophxt<Mi crooksu Benson) L Benson Amcr
2.
(J B^
J
ART/
EX) T.x\s Piso Co Eiinkhn Mts mi
El Trins
V( jct
I 1 1
N
W)
'» (M 2^
and Gitewav S >3 106 southern exposuie of
26 Aug 19-^8 }2(raLX) Hudspeth Co hme
WorthiN^t,,?
-x
Williams, Notes on Haplophyton 47
3
Quitman Apr
Mts, 17 1976,
basin in the
W
among
mouth of narrow canyon draining S slope of the Chinatis, igneous boulders,
4800 5857b (TEX).
14 Jun 1977, Butterwick
ft,
km ENE
MEXICO.
Chihuahua: ca 20 of Ciudad Jimenez, in limestone arroyo in can-
NW
yon of summit of Sierra de Chupaderos (27°12'N; 104°43'W), 5100 ft, 2 Oct 1973,
NW
SW
Henrickson 13765 (TEX); ca 31 mi of Julimes in facing canyon above Rancho El
4400
Recuerdo de Carrasco (28°47'N; 105°09'W), 15 Sep 1973, Henrickson
in Sierra ft,
W
km
km
12941 (TEX); 46 of Ojinaga on hwy to Chihuahua City and S 2 to the deep
104°48'W), 1000 m, 20 Oct
canyon of Rio Conchos in the Sierra de Pegiiis (29°32'N;
NNE
1972, Chiang 9757 (TEX); S slope and top of Sierra del Roque, of Julimes
ei al.
approached from Mina Las Playas via Rancho El Saucito (28°39'N; 105°18'W), 19 Jun
973, M.C.Johnston 11386B (TEX). Coahuila: S side of Cerro San Jose de las Piedras
et al.
1
km
E
(28°42'N;102°51'W), 1000-1400 m, 8 Jun 1972, C^/^«^e^^/, 7355 (TEX); 0.5 of
MatrimonioViejo,E end of limestone ridge (27°08'N; 103°07'W), 1125 m, 2 Sep 1972,
800-2000 17-25
Chiang 9122 (TEX). Sonora: Sierra Bojihuacame SE of Obregon, ft,
mi
Oct 1954, Gentry 14482 (TEX); talus slope of low basaltic hill, 15 S of La Palma,
Mayo
between La Palma and Guaymas, 2 Sep 94 Wiggins 226 (TEX); Rio area, hill with
1
1 ,
NW
SSW
microwave tower, 9 mi of Alamos and 4 mi of Piedras Verde (27°07'N;
km NE
Canon
109°02'W), 500-700 m, 6 Sep 1989, Sanders 9435 (TEX); de Nacapules, 6
of Bahia San Carlos, deep riparian canyon, 19 Oct 1984, Felger 84-122 (TEX).
monogeneric
oi Haplophyton, Pichon (1950) erected the
In his classification
He
with
subtribe, Haplophytinae. discussed its affiliation the
remarking
Catharanthinae, but ended by questioning the relationship, that
"the lignification of the branches [in Haplophyton'] seems to be rather rapid
common
view with
and the plant has nothing in in this point of the
Leeuwenberg subsequently
Catharanthinae" (Pichon 1950; 6 (1994)
p. 1 1).
positioned Haplophyton with Alstoma R. Br. in the Alstoniinae, near the
two
Catharanthinae, on the basis that both genera are the only taxa in the
Plumerioideae to possess hairy seeds. Seed pubescence in the Alstoniinae,
and
however, heteromorphic. Haplophyton has seeds with both apical basal
is
membranous
coma, while Alstonia has seeds with either ciliations (sect.
coma Monuraspermum)
Tonduzia) or with both apical and basal (sect.
(Monachino
1949).
The dextrorse aestivation found in Haplophyton, suggests a close rela-
which Indeed,
tionship with Alstonia, also displays dextrorse aestivation.
may
Haplophyton cimkidum be related to the Oceanic species of Alstonia
and
Monuraspermum), which have flowers with dextrorse aestivation
(sect.
coma (Monachino how-
with both and 1949). Presently,
seeds apical basal
ever, not known whether or not dextrorse aestivation in Haplophyton is
it is
ancestral or derived.
Alstonia the only other genus in the tribe Plumerieae, besides the gen-
is
of the Catharanthinae (excluding \tnca (Lawrence 1959)), to retain a
era
membranous appendage stigma throughout ontog-
reflexed basal to the its
Woodson
eny (Pichon 1950). (1928) noted, however, that a swollen region
474
SiDA 1995
16(3)
below
occurs the stigmas oi Haplophyton. In his view, this character prob-
ably represents a primitive stage of the more complex appendage found
in
the Catharanthinae. Preliminary studies of Haplophyton herbarium
speci-
mens, by the author, reveal that the heads of immature buds have
style
a
membranous
skirt below them. This feature, however, was not present
in
mature
(opened) flowers in either the or herbarium specimens.
field
Pichon
(1950) also suggested an between Haplophyton and
affiliation
two
Anechites, stating that the genera are unique in the family by having
with
hairs multicellular bases (Fallen 1983). Pichon, however, placed each
of these genera in monotypic subtribes, suggesting that their relationships
were unresolved. was subsequently
Anechites positioned in the subtribe
Condylocarpinae
of the subfamily Plumerioideae (Leeuwenberg
1994).
Haplophyton from Monuraspermum
differs Alstonia being
sect. in a
suffruticose herb (vs. tree), having alternate (vs. whorled) leaves, hairs with
multicellular bases simple), no nectary an annular
(vs. (vs. nectary), soli-
New
tary flowers (vs. cymous), and its restriction to the World (vs. Oceania).
These
differences suggest that Haplophyton not related to Ahtonia. Until
is
further evidence presented, the most suitable and conservative course
is
favors placing Haplophyton in the monogeneric subtribe Haplophytinae
(Pichon 1950) of the tribe Plumerieae, and not with Ahtoma the
in
Alstoniinae (Leeuwenberg 1994) whose resemblance to Haplophyton
at
is
present questionable.
am
grateful to Carol Todzia, James Henrickson, and the two anony-
I
mous my
reviewers for their attentive editing of the manuscript, and
to
companion
work
Jennifer Forrest for translating Pichon's from the original
French. B.L. Turner provided helpful comments. Cooperation from and
F,
TEX-LL much my
was
acknowledge
also appreciated. Finally, friend Jon
I
Plum, having accompany me
for the tenacity to to Chiapas, despite the
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