FOLIA TAXONOMICA 16. DILKEA (PASSIFLORACEAE) 1. EPKIA, A NEW SUBGENUS AND FIVE NEW SPECIES FROM WESTERN AMAZONIA AND THE GUIANAS

2009·15.1 MB·English

by  Christian Feuillet

TAXONOMICA
FOLIA DILKEA (PASSIFLORACEAE)
EPKIA,
16.
1.
NEW NEW
AND
A
SUBGENUS
FIVE SPECIES
AND
FROM WESTERN AMAZONIA
THE
GUIANAS
Feu
Christian
let
il
MRC-166
Department of
Botany,
i a Dilkea (Passifloraceae) e tpres nteee le sous-genre Epkia et trois especes nouv
cuneata, D. ovalis D. ue deux especes des Guyanes: Dilkea lect
et ti
tlfaaeaau Lpreentee
RESUMEN
ntroduccionalgenero Dilke a (Pass florae ae) y se describe el nuevo subgenera Epkia
yem
nasco aelpr mer
subgenera Epfeiaposeenra entrenudo escamas de
tras las la
was named by honor Wentworth made
Dilkea Masters (1871) his friend Charles Dilke, (1810-1869),
to late
II
by Queen was and He
baronet Victoria in 1862. Dilke a patron of arts sciences. died in St Petersburg while
representing England at the horticultural exhibition held there. His father, C.W. Dilke, (1789-1864), was
I
founder
a of the Gardener's Chronicle.
with genus The can be from
Dilkea, 7 species, a of tribe Passifloreae. species easily separated the other
is
American on
genera of the based the following characters flowers with 4 4 8 stamens,
tribe suite: sepals, petals,
mm
4 carpels and styles, stamens inserted at the bottom of the tube or halfway up the short, 0-2 long an-
drogynophore, and nor ornamented Holm-Nielsen
seeds that are neither flattened (Masters 1872; Killip 1938;
&
MacDougal Panama
1988; 2003; Feuillet 2007). Dilkea has a large lowland distribution in
et Tillett
al.
and South America, from Colombia Mato Grosso and from Peru Amapa, but has been
to to collected.
little
Dilkea SOC
Mast., Trans. Linn. 27:627. 1871.
Type species: D. retusa Mast., Trans. Linn. Soc. 27:628. 1871. (for typification,
Shrubs, small Leaves always simple and unlobed, margin with
trees, or lianas. entire; petiole a basal pulvi-
Flowers operculum and limen androgynophore than wide
nus. white; sepals petals lacking; shorter or
4;
4;
mm
lacking; stamens inserted on the bottom of the floral cup or on a 1-2 long androgynophore; ovary
8,
mm
sessile or usually on a short 1-2 long gynophore, styles 4, united at base. Fruits yellow to orange, bac-
ovoid rounded with acumen,
cate, or globose, apically or a conical pericarp
c<
Journal of the Botanical Research Institute of Texas 3(2)
594
and
bean-shaped or not flattened, both ends similar, sarcotesta thin, neither sclerified
few, slightly straight,
There morphology within Dilkea; thus, one cannot separate species in the
a great similarity in floral
is
am
and commented:
on described Dilkea
genus based characteristics alone. Killip (1938) parviflora "I
floral
show
think further study and collecting will that the four last species of the present treat-
inclined that
to
number
ment should be merged in one." Part of the problem, as suggested by Killip, resided in the small of
synonymizing sampling, appears that
but instead of getting a species as a result of better it
collections,
field
new
increased sampling instead revealing species.
is
nine species have been described within Dilkea. Masters (1871) published D. acuminata
Historically,
new
new Masters de-
within genus. (1872)
and from Spruce's collections as species this Later,
D. retusa
Bora
only an by Wallis, in his treatment of the Passifloraceae for
scribed D. Mast., after illustration
wallisii
by
by Rodrigues
Few Barbosa D.
have been published since then: D.johannesii (1888), ulei
taxa
Brasiliensis.
Hoehne
by
Harms (nom. nud.) by Glaziou (1909), D.johannesii var. parvijolia (1915), D.
(1906), D. glaziovii
more
and by Cervi For
by by Steyermark D. margaritae (1991).
D. (1968),
magnifica
Killip (1938),
parviflora
synonym Mast, ex Glaz. (nom.
been Dilkea
than 70 years, D. has treated as a of D. johannesii. glaziovii
ulei
Thus, present
was Mitostemma Mast, was considered to be in the right genus. at
nud.) ignored, as glaziovii
seven species and one variety are circumscribed within Dilkea.
synonymy
and
Most Holm-Nielsen (1988) placed D. acuminata, D. magnifica, D. wallisii in
recently, et
al.
and
on
and separate species the basis of petiole, pedicel,
keeping D.johannesii D. parviflora as
of D. retusa,
on
and based
gynophore They had not seen the type of D. therefore their identification Kil-
parviflora
sizes.
new
description. In the material cited from Ecuador and the illustration mostly belong to a species
fact,
lip's
below
described as D. cuneata.
an unexpected
Smithsonian herbarium (US) revealed
Recent of Dilkea collections to the Institution
gifts
was confirmed
and from This
morphology from the Loreto region in Peru the Guianas.
diversity of vegetative
in February 2009 by a visit to the herbarium collections at the Missouri Botanical Garden, especially rich in
woody monopodial growth
continuous
from western Amazonia. Several species are vines that exhibit
Dilkea
mean but deciduous they do not find a support.
and climb by the of strong tendrils apically 3-fid early if
Another group including the
They what considered here be Dilkea subg. Dilkea. of species,
represent to
is
and
new described below, have similar flower and fruit morphology, but are small trees exhibit a
species
five
The stems have strongly modular growth;
unique "modular" growth pattern that rare in the Passifloraceae.
is
and
bud
with then long internode, apically a
modules have the base very short internodes scales, a
the
at
swelling with short internodes, leaves, and inflorescences and ramifications. In most cases, the terminal
meristem aborts and the growth continues through l(-3) subterminal vegetative buds. This results in the
modular growth
rhythmic
appearance with one stem and of leaves, evidence of or
overall of a treelet tiers
and sometimes few modular branches 1-2 meters above ground.
needed a
ascertain
studies are to this)
(field
The standing and do not have tendrils.
plants are
self
and
In the below, the world "lacking" means "not present in the taxon" the expression "not seen"
text
Subgenus Epkia
Epkia
Dilkea subg. Feuillet, subg. nov. Dilkea cuneata Feuillet (see below).
Type:
The subgenus Epkia are mostly small trees (or shrubs or scandent shrubs). In each branch, the first
species of
much and no
node numerous bud longer than the following internodes there is
above the acicular scales is
become branches higher rami-
shape and morphology. The long internodes shorter in the of
transition in leaf
growth
The in a terminal cluster on the club-shaped apical part of every
order. large leaves are alternate
fication
by and morphology.
general
unit. The flowers are similar to those of the other species of Dilkea their color
A new new
subgenus and
Feuillet, five species of Dilkea 595
m
Collection labels usually say "tree" or "small tree" with noted sizes from <l-8 but are occasion-
tall,
ally labeled as "shrub." think this is indicative of a small size rather than a clumping habit of the plants.
I
m
From
cm
pictures taken in the wild, the young plants 0.5-1.5 have main stem about
tall a 1 in diameter,
A
many
showing
few
leaves in a tiers. 1.5-m tall plant of Dilkea cuneata photographed in Peru has 4
tiers
of 7-20 large leaves. The lower is at ground level and the upper one at the apex of the plant. The picture
is
shows whole and
the plant not possible decide the stem has monopodial and rhythmic growth
is to a or
it if
a sympodial, modular growth. The branches, as demonstrated by herbarium specimens, modular and
are
branched Each module
little constructions. exhibits a unique morphology. At the base are usually more
bud
than by and
10 acicular scales followed a long internode the apical part with short internodes bearing
and
the leaves axillary flowers or short inflorescences.
name
The
of subgenus Epkia honors through his Ellsworth Paine (1890-1968) author
initials Killip of
The American two-volume 366
Species Passifloraceae (1938), a masterpiece covering species.
of
KEY TO THE SUBGENERA OF
DILKEA
internodes; stamens inserted at the bottom of the hypanthium. Dilkea subg. Dilkea
much
Small trees or shrubs or scandent shrubs; without tendrils; internode longer than the following
first
1
.
mm
stamens on 1-2
internodes; inserted a long androgynophore. Dilkea subg. Epkia
species from western Amazonia and two from the Guianas here described subgenus
are in
Epkia.
of Dilkea are undescribed.
is still
Dilkea cuneata
1. Feuillet, sp. nov. (Fig.
1). i
m
1-5 monocaulous 6-20
Treelets or branched. Plant glabrous throughout. Stems with long internode
a
tall,
mm
cm cm
cm
long (2.5 in Gentry et 54258), 0.4-0.6 thick, followed by short ones 0.2-0.5 long. Tendrils
al.
lacking. Stipules not seen. Leaves in terminal clusters; petiole reduced the pulvinus 0.8-1.5 cm; blade
to
28-53
narrowly cuneate to cuneate, x 10-13.5 cm, short long attenuate and then abruptly ending
to base,
at
mm
apex acute rounded and acuminate, acumen 5-15 margin drying
to long, olive-green
entire, adaxially,
brown
yellowish midrib and on both and when 12-25 main
abaxially, veins raised surfaces yellowish
dry,
mm
on
each
veins side of the midrib. Inflorescences subterminal; pedicels about 5 long, elongating 7-10
mm
cm
under hypanthium
the bracts not seen. Flowers scented; 0.5-1.8 funnel-shaped, 0.4-0.8
fruit; long,
cm
diam. at throat; sepals white, oblong, 2.5-3.5 x 0.7-1 cm, obtuse to acute apex, margin;
4, at ciliate at
petals white, oblong, 2.5-3.5 x 0.5-0.7 cm, obtuse acute apex, united thinner than
to base,
4, at at sepals;
cm
corona
white, in 2 rows, outer one of free straight filaments, 1-3 long, seldom curled apex, inner one
at
mm
membranous,
margin, androgynophore 1-2
laciniate at laciniae floccose; long; stamens filaments
8,
mm
cm mm,
white, 3 united base anthers sulphur 8
ca. long, at for yellow, long, dorsifixed near base;
1
mm mm
cm
5-6
gynophore 1-2
ovary
long; ovoid, long, styles ca. 3.5 long, united for 1/3-1/2 of their
4,
cm
length, stigmas capitate, sulphur yellow. Fruits spherical or slightly ovoid, apiculate, 2.5-3 long, yellow,
mm
pericarp coriaceous, about 0.5 thick, glabrous; seeds few, elongate, 1.5-1.7 x 0.7-0.8 cm, brown.
—
known
Amazonian
and
Distribution ecology. Dilkea cuneata a western species from Ecuador (Morona-
is
m
and Peru and primary on 100-700
Santiago) (Junin, Loreto, Pasco), in forest terra firme, at of elevation.
documented blooming and November, and
It is in February, June, July, fruiting in January to April, June,
and
October.
July,
Several specimens lacking flowers have been misidentified as belonging to the Theophrastaceae Clavija
Journal of the Botanical Research Institute of Texas 3(2)
&
Smith 26730
Dilkea cuneata, holotype Killip (US).
Fig. 1
.
A new subgenus and new
Feuillet, five species of Dilkea 597
Hook. by
julgens (including Killip in 1931), Clavija lehmannii Mez, Clavijapoeppigii Mez, and
F. to Passiflora L.,
member
Some
even as a of the Ranales. specimens have been identified as Dilkea (Holm-Nielsen
parviflora
and
et 1988) Dilkea wallisii (Killip 1938) although those two Dilkea species are lianas. Nevertheless, D.
al.
new
from western Amazonia. some
cuneata clearly represents a species Interestingly, D. cuneata has of the
cm
largest leaves and of the smallest flowers in the genus. The growth units have 15-25 long internode
a
bud
between the scales and the leaves and the leaves are crowded toward the apex with internodes shorter
all
than or equal to twice the diameter of the petioles. The congested inflorescence the end of a branch not
at
is
but
terminal, in fact consists of 2 or 3 axillary, subsessile, pauciflowered inflorescences. Aestivation of the
two
outer sepals valvate, this pair surrounding the two inner sepals.
is
Etymology.—The
epithet "cuneata" refers to the leaf blades basal half that are long triangular.
m
with branches sometimes up Whole
Treelets passive climbers, 3 plant glabrous. Stems with long
to a
tall.
cm
internode 1.3-5 long, followed by short ones about as long as once their diameter or Vegetative
less.
mm
mm
bud bud 3-5
above
ca. 2 the leaf axils, scales acicular to narrow-triangular long. Tendrils lacking.
Stipules not seen. Leaves in terminal clusters; petiole usually reduced to the pulvinus, occasionally up to 0.6
cm
narrow 8-17 2-5
long, glands not seen; blade oblanceolate, cuneate, x cm, long apex
attenuate, acute,
long acuminate, margin midrib and on both and when 8-12
entire, veins raised surfaces yellowish dry,
main
on
each
veins side of the midrib. Inflorescences subterminal; pedicels not seen. Flowers not seen
(see
mm
gynophore 1-2 under
below); long the united only
fruit; styles 4, at base. Fruits spherical, apiculate,
mm
3-3.5 2.5-3 cm, documented,
x color not pericarp coriaceous, about 0.5 thick; developed seeds not
seen, ovules 2 per placenta.
flat,
—
known
Distribution and ecology. Dilkea lecta from the Lely Mountains in eastern Surinam and from
is
and
central northwestern French Guiana, and Brazil (Matto Grosso). The ecology different between the four
is
collections. Loubry 2422 was collected in primary forest on white sand, on the coastal plain, low elevation
at
map
(according to the the whole area below 50 m); Larpin 1058 from low forest island on the northern
is is
&
340 m; was
slope of a granite outcrop (inselberg) at Mori Bolten 8541 collected between 2 plateaus of a
&
mountain range between 500 and 700 m, and 13648 from middle high
Granville Crozier ridge
is to forests
on and conglomerate 400 m. The two French Guiana low amount
schist, quartzite, at localities in share a of
available water due to the texture of the white sand without visible amount of humus causing excellent
soil,
maximum
and and
drainage toward undersurface one and dark
fast light reflection leaf in case the granite
slope preventing water storage and favoring evaporation.
Dilkea distinguished from D. cuneata by leaves three times smaller and by the internode modu-
lecta is its
Journal of the Botanical Research Institute of Texas 3(2)
Loubry2422
Ditkea holotype (US).
2. lecta,
Fig.
A new new
subgenus and
Feuillet, five species of Dilkea 599
While
larity so characteristic of subg. Epkia. this branching pattern less obvious than in the other species
is
&
subgenus main
of Epkia, visible. Plants of Dilkea are shrubs likely with one trunk
is still lecta (Granville
it
&
Crozier 13648 and Loubry 2422), yet able to passively climb on small trees (Larpin 1058 and Mori Bolten
85411
Hoehne
and
(1915) gave a brief description a diagnostic illustration of Dilkea johannesii
var. parvijolia.
synonym
most
This likely a of D. Based on Hoechne's description one can
is lecta. infer the following: leaf
cm
blades convex between veins, with small red dots underneath, pedicels more than
slightly long, flower
1
hypanthium
with asymmetric and
slightly at base), sepals petals of similar length, corona in 2 rows,
at least
the inner one inserted low in the hypanthium, outer row of filaments exerted (more than
ligulate, slightly so
membranous
the stigmas), inner one base, filamentose apex, filamentose segments bare and
at at basal 1/3
at
upper
stamens than on
2/3 papillose; shorter the petals, inserted a short androgynophore; short gynophore;
&
wide
ovary elliptic, styles united for 2/3 of their length (unlike Mori Bolten 8541), stigmas spherical. Thus,
my
on
based
these features, this species within current circumscription of D.
fits lecta.
Etymology.—The
specific epithet "lecta," Latin for gathered, refers to the leaves grouped at the apex of
the branches.
Dilkea nov
OValis
3. Feuillet, sp. (Fig. 3). PERU. Loreto: Maynas, Las Amazonas, 72°33W, 120-130 m, 25 Mar
Type: 3°25'S,
m
4-6
Shrub cm
or small plant glabrous throughout. Stems with long internode 3.5-12
tree, tall; a long,
cm
by
followed short ones 0.1-0.3 long. Tendrils lacking. Stipules not seen. Leaves in terminal
clusters;
cm
0.5-2.3
petiole long, pulvinate for less than half length, glands not seen; lamina
its elliptic to slightly
obovate, 7.5-25 x 3.5-10 cm, widely acute and then short cuneate apex rounded with acumen
at base,
cm
when
margin
0.6-1.2 long, minutely midrib and on both 6-12
entire, frilled dry, veins raised surfaces,
mm
main
veins each side of the midrib. Inflorescences subterminal; pedicels under 7-9
fruit long; bracts
mm
not seen. Flowers not seen; gynophore under about
fruit 1 long. Fruits spherical or slightly ovoid, api-
mm
cm
about
culate, 2.5 diam., yellow, pericarp coriaceous, about 0.5 thick, glabrous; seeds few, elongate,
1.8-2 x 0.8-0.9 cm, brown.
known
Distribution and ecology.—The three collections have been made in northeastern Peru, Region
of Loreto, Province of Maynas, at 140-180 m. was in flower buds in July and fruiting in March.
It
Dilkea ovalis is easily set apart from other species in subgenus Epkia by leaf shape. In D. the
ovalis,
laminas are wide elliptic rather than cuneate or narrow elliptic. The pulvinus a third to half the petiole
is
total length. In the other long-petioled species, D. vanessae, the pulvinus represents less than the of the
fifth
length.
total
—
named
Etymology. This species is ovalis because of the oval shape of leaf blades.
its
PERU. 47"W
Maynas, 73°25
Paratype: Loreto: Iquitos, Allpahuayo-IIAP, , 3°57'19"S, 140 m, 26 Apr 1997, Vdsquez,
R. O.
St., Phillips,
)
Jammillo 17188 (MO).
Dilkea
4. tillettii Feuillet, Sp. nov. (Fig. 4). Type: PERU. Loreto: Florida, Rio Putumayo mouth of Rio Zubineta, 200 m,
at ca.
Mar-Apr MO,
Klug2100
1931, US;
G. (holotype:
isotypes:
fl., S).
= unknown 986 on
[Dilkea grandifohai tied. handwriting 1 Gentry et at 3 7992 (F) Tillet" and Tilletts hand-
,
m
1-5 monocaulous
branched.
* or
tall,
I
Texas
Journal of the Botanical Research Institute of 3(2)
I
Grandez 2291 (MO).
holotype et
mils, al.
new new
A subgenus and
Feuillet, five species of Dilkea
x
a.
.
Klug2100
4. Dilkea holotype (US).
Fig. tillettii,
Journal of the Botanical Research Institute of Texas 3(2)
602
cm
cm by ones 0.2-0.5
37992 followed short
>12.5 long (Gentry
node, the only partially visible et (F)),
al.
cm
reduced
not Leaves in terminal clusters; petiole 1.5-3 long, usually
ong. Tendrils lacking. Stipules seen.
35-43 12-15 cm, long triangular
x basal half
pulvinus, glands not seen; lamina long oblanceolate,
o the
cm
acumen up
with an
narrowing the short pulvinate petiole, rounded at apex, blunt or to 1.5
regularly to
margin midrib and veins raised on both surfaces and drying more or less the same color as
entire,
cm
8-12 main subterminal; pedicels 0.5
veins each side of the midrib. Inflorescences
the
of leaves,
st
cm with
cm from not Flowers white, a scent of
under joined 0.2-0.3 the base; bracts seen.
the
long,
fruit,
1
mm
and hypanthium during
hypanthium 17-19 perianth separating getting shorter
long,
(Barrier
lilac 84);
mm
mm
white
and 20-35 about wide, two outer sepals green outside,
anthesis; sepals petals long, 5
e
two
on both corona white in
two inner sepals white surfaces, aestivation valvate;
aestivation valvate,
side,
cm row wavy,
ws 3.5-4 long, inserted near base of hypanthium, tubular at base, outer filaments slightly
and stamens
branched uppermost cm, segments very thin tightly curled;
heavily in the lateral
ner series 1
mm,
cm
llx base 2
bottom on hypanthium, filaments about 3 long, anther apiculate,
>erted in the the 1
mm
mm
mm
3.5-4
wider
apex,
gynophore about 2 long; ovary 5 long, tapering at base, at styles
sagitate;
cm
about
cm 2.5-3 diam., apiculate, yellow, pericarp coriaceous, 0.5
stigma Fruits globose,
long, capitate.
mm
cm, brown.
x
glabrous; seeds few, elongate, 1.8 0.8
thick,
&
known Ecuador
Colombia (Amazonas
from Caqueta),
and ecology.—Dilhea
Distribution is
tillettii
&
&
between 120-700 m. was blooming in
Napo and Peru (Loreto Pasco)
(Morona-Santiago, Pastaza), It
and November.
September
January and March-April, and fruiting in June to
and
has obovate leaves that are cuneate in the basal half unlike D. lecta D. ovalis. It is
Dilkea
tillettii
by shape and the shape of the
the of the petioles
from the other large-leaved species, D. cuneata,
distinct
more than length the does not always
Although the pulvinus constitutes half the of petiole, it
base.
leaf
narrow shape,
encompass The base the lamina the continuation of the triangular leaf
the entire petiole. of is
more reduced
has or
neither suddenly ending as in D. cuneata nor decurrent. Dilkea lecta also petioles less
most
the pulvinus, but laminas are at half the size of those in D. tillettii.
to its
MYF), and
named honor botanist in Caracas a specialist
Etymology.— Dilkea to Steve Tillett, (at
is
tillettii
new
who
was
on the Passifloraceae recognized this a species.
Caquci
16039 (MO).
1991, Pipoly
fr.,
J.J.
&
Asanza 31937 (AAU);
Brandbyge
E.
J.
5. Dilkea vanessae Feuillet, sp. nOV. (Fig. 5). Type: FRENCH GUIANA. Pic Matecho, camp near the summit, 500 m, 14-15
m
cm
2-20 by
followed short
0.8-2 monocaulous or branched. Stems with a long internode long,
Treelet
tall,
cm
cm 2-2.5
long,
ones 0.2-0.3 Tendrils lacking. Stipules not seen. Leaves in terminal clusters; petiole
long.
cm 19-21 7-8 cm,
x basal
including the pulvinus 0.3-0.4 long, glands not seen; blade oblanceolate, cuneate,
rounded
apex
and then with very narrow wing along the petiole until the swollen base,
half triangular a
when
brown
and margin midrib and veins raised on both surfaces and light dry,
long-acuminate,
entire,
mm
bud,
7-9 main veins on each side of the midrib. Inflorescences subterminal; pedicels ca. 6 long in joint