Three new endemic specis of Cremastosperma (Annonaceae) from the Rio Marañon basin, Amazonas, Peru

2004·5.9 MB·English

by  M D Pirie

Amaldoa
7-20 .2004
11(2):
Three new endemic Cremastosperma (Annonaceae)
species of
Maranon Amazonas, Peru
from Rio
the basin,
MICHAEL
D.PIRIE
Herbarium Nederland,
Nationaal
Utrecht branch,
Universiteit
CS
3584
Heidelberglaan Utrecht,
2,
The Netherlands
[email protected]
CRUZ
MARIO
ZAPATA
Museo
de Historia Natural,
Antenor Orrego
Universidad Privada
Trujillo-PERU
@
mzapatac upao. edu.pe
Abstract
Subsequent a expedition the Peruvian department of Amazonas, and in
to field to
new Cremastosperma (Annonaceae),
advance of a revision of the genus, three species of
endemic to the Rio Marafion basin in Peru, are described here: C. bullatum Pirie, C.
&
yamayakatense and cenepense Zapata.
Pirie C. Pirie
Resumen
Como Ama-
resultado de los trabajos de campo realizados en el Departamento de
zonas (Peru); y en avance a la revision del genero, describimos tres nuevas especies de
C
Cremastosperma (Annonaceae): bullatum yamayakatense
C. Pirie, Pirie y C.
&
cenepense endemicas de cuenca Maranon.
Pirie Zapata, la del rio
The genus Cremastosperma was described by Fries 1930, and phylogenetic analyses
DNA
using sequence data suggest that species comprise a monophyletic group Pirie
its
&
al., in press). It can be distinguished from other Neotropical Annonaceae by its raised
midrib with a unique longitudinal groove. Although the time of writing only 18 species
at
of Cremastosperma are recognised, the results of ongoing systematic research suggest
number may The
that the total of distinguishable species in fact exceed 30. distribution of
species of Cremastosperma can be summarised according to Gentry 1982) as 'Andean
(
centred' are largely absent from Amazonia, but diverse in middle elevation cloud
thiey
:
forests and in the lowlands near the base of the Andean mountain chain. These regions
coincide with two of the twenty-five areas worldwide designated as biodiversity hotspots
&
by Myers Choco/DarienAVestem Ecuador and An-
(2000): the region the tropical
al.
The Andes
des. area of the tropical in Peru particularly rich in species of
is
Cremastosperma, housing 12 of the 18 currendy recognised species.
new
Three species of Cremastosperma from Peru are described here: C. bullatum
C
&
yamayakatense and cenepense Cremastosperma
Pirie, Pirie, C. Pirie Zapata.
bullatum and C. yamayakatense were collected by the authors during a field expedition
in northern Peru at the end of 2003. Both were found in the area of the community of
Yamayakat in Bagua, Amazonas, from where a further species of Cremastosperma, C.
peruvianum by and by
(described Fries, 934, distinguished in particular long, relatively
1 its
narrow leaves with cordate base, and long stipes), has also been collected. These three
C
new known
and cenepense (the third species described here) are only from herbarium
Maranon
specimens collected in the Rio basin in Peru. This region appears not only to be
unusually rich in often narrowly endemic species of Annonaceae, but also until recently to
have been relatively overlooked in terms of both collections and taxonomic work. Collections
of the genera Unonopsis, Rollinia, and Guatteria made during the same expedition
number
appear to represent a of as yet undescribed species Maas, pers. com.).
(P.
-
Cremastosperma bullatum 3-5
Pirie spec. nov. Fig.
/. 2,
Typus: Pirie, M.D.; Zapata C, M.; Apanu N., R. 71 (holo: U; iso AAU,
AMAZ, CUZ, HAO, MO, MOL, USM,
NY, HUT, US,
E, K,
F,
WU), Peru. Amazonas: Bagua, Imaza, community
District
Yamayakat, Putuim, 22 Nov 2003
to
trail
(fl).
margine indumento
Foliis bullatis ciliata, e pilis longis composite, pedicellis longis,
petalis interioribus petala exteriora superantibus distinctum
m
TVee 2- 10 young twigs and densely covered with mainly
tall; petioles erect golden
mm mm mm
up
hairs to long. Leaves: petioles 3-7 long, 2.5-3 diam.; lamina or
1 elliptic
cm
narrowly so to slightly obovate, 17-28 by 6-1 (leaf index 2.4-3.5), chartaceous, mid
1
brown, occasionally slightly grey above (immature leaves drying black), glabrous or sparsely
mm
covered with mainly erect golden hairs up to long on upper surface, densely so on
1
all veins below and on edge of lamina, base rounded to subcordate, apex acuminate
mm
(acumen 5-20
long), extreme tip obtuse, primary, secondary, and tertiary veins sunken
mm
in depressions in leaf surface, primary vein raised along entire leaf length, .5-2 wide
1
mm
at widest point, densely covered with mainly erect golden hairs up to long above
1
and below, secondary veins 15-20 (intersecondary veins distance between from 6
rare),
mm mm
at the base to 16 closer to the apex, angles with primary vein consistently around
60-70°, occasionally branching, forming distinct loops, smallest distance between loops
mm,
and margin 1-1.5 tertiary veins largely percurrent. Inflorescences of single,
successively produced, pendulous flowers, axillary on leafy branches, on leafless branches
woody
and produced from the main trunk (then extending from a structure comprising the
mm mm
remnants of old inflorescences); short axillary shoot, 17-20 long, 1-1.5 diam. (in
mm mm mm mm
flower), 18-25 long, c. 1.5 diam. (in fruit); pedicels 100-140 long, c.
1
mm mm mm
diam. at the base, 2 diam. at the apex (in flower), c. 1 .5 diam. at the base, 2
apex and
diam. at the (in fruit); short axillary shoot pedicel sparsely to rather densely
mm
covered with mainly golden up lower
erect hairs to long, single bract, elliptic to
1
mm mm
ovate, 2.5 long, wide, acute, persistent or partially caducous, densely covered
1
mm
with mainly erect golden hairs up to long; upper bract within central third of pedicel
1
mm mm
length, elliptic to ovate, 2-3 long, 1-2 wide, acute, densely covered with mainly
mm
erect golden hairs up to long; flower buds depressed ovoid developing to conical,
1
brown
green in vivo, in sicco; flowers green, maturing to yellow with a basal orange patch
on the outside of the outer petals in vivo, golden brown in sicco, outer sides and apical
portion of the inner sides of petals and outer sides of sepals densely covered with appressed
mm
golden hairs up to long, inner sides otherwise glabrous; sepals basally connate,
1
mm,
mm,
deltate, 5-7 by c. 6 acute, caduceus; outer petals broadly ovate, c. 18 by c. 15
mm
inner petals ovate, concave, 25 by c.l2 wide; receptacle ovoid; androecium 5
c.
c.
mm mm
diam., stamens 1-1 .5 long, connective appendage of inconsistent and irregular
mm
shape, c. 0.5 wide, glabrous. Monocarps 8-10, dark brown in sicco, ellipsoid, slightly
mm mm
assymmetrical, c. 5 long, diam., apicule (where visible) excentric; stipes 14-
1 1 1
mm mm mm
16 long, c. 1.5 in diam; fruiting receptacle ovoid, 5-6 in diam., monocarps,
mm
and
stipes receptacle sparsely to rather densely covered with erect golden hairs
0.
1
mm mm
many
Seeds orange-brown
long. ellipsoid, with shallow c.l3 long, 10
pits, c. in
diam., raphe raised, encircling seed longitudinally,
i
m
and
Habitat ecology: Trees or shrubs from 2 have been
collected primary
tall in
flower February and November, November and
forest in in in fruit in June.
Other specimens examined:
PERU: Amazonas, Bagua: 20 Jan 1996
Jaramillo, N.; et
(fl, fr), al.
942 (MO, 25 Jan 1996 972
U); Jaramillo, N.; et (HUT,
(fr), al.
MO,
community
Imaza, Yamayakat,
U); District to Putuim,
trail
22 Nov 2003 66 HAO, USM);
M.D.; et U,
(st), Pirie, al. (F, (fr),
M.D.; 68 (HAO, US, USM); Imaza,
et U, District
Pirie,
al.
community Putuim, path leading west along Quebrada
Shimutaz, 25 Nov 2003 M.D.; et 94 (AMAZ, CUZ,
Pirie, al.
(fl),
HAO, HUT, MO, MOL, USM); Imaza,
K, NY, U, District
community Aguaruna de Putuim, 20 Jun 1996 Rodriguez
(fr),
community
1152 Imaza, Yamayal<at,
R., E.; et (U); District
al.
15 Nov 1997 Vasquez, 24891
R.; et (U).
al.
(fr, fl),
Notes Cremastosperma bullatum can be distinguished from other species
easily
all
:
of Cremastosperma by any one of the number of unique and striking characteristics
it
when
displays. The leaf blade has a blistered or bubbled appearance, both in the field and
pressed, which due to the deeply sunken nature of the primary, secondary, and tertiary
is
venation. The indument present on many of parts far longer than in any other species
its is
in the genus, and, also uniquely in the genus, densely inserted in a halo-like formation
around the leaf margin. Other notable characteristics are the unusually long pedicel, the
orange colouring of the base of the outer petals of mature flowers, the inner petals
considerably longer than the outer petals, and the rounded to subcordate shape of the leaf
Etymology The
bubbled appearance of
specific epithet reflects the the leaves.
:
Cremastosperma yamayakatense -Fig. 6-8
Pirie spec. nov.
2. 2,
Typus: Pirie, M.D.; Zapata C, M.; Apanu N., R. 57 (holo U; iso CUZ,
HAO, HUT, MO, USM), Amazonas: Bagua,
NY, Peru.
K,
Imaza, community Yamayakat, Putuim, 22 Nov
District to
trail
A
C. gracilipes pedicellis brevioribus, alabastris glabris ante anthesin inapertis
differt.
A
cenepense monocarpiis longioribus
C. stipitibus differt.
m
Tree 1.5-8 (-20) young twigs and petioles shallowly grooved, glabrous.
tall;
mm
Leaves: petioles 5-10 long, 1-5 mmdiam.; lamina 1-24 by 3.5-8
elliptic, 1 (-38) (-13)
cm
(leaf index 2.4-3.4), chartaceous, olive-grey green above, light brown below, glabrous
mm
on both base apex acuminate acuminate (acumen 10-25
sides, acute, shortly to long),
extreme tip rounded, primary vein raised over entire leaf length, grooved for 25-30%
first
mm
from base, 1-4 wide at widest point, glabrous, secondary veins 8-10 (-14),
mm
intersecondary veins occasional, distance between from 5-10 at the base to 10-30
mm
closer to the apex, angles with primary vein from 70-80°, the angle thereafter
10
forming
decreasing and subsequently increasing towards the leaf margin, not branching,
mm,
distinct loops, smallest distance between loops and margin 2-6 tertiary veins largely
some of successively produced, flowers,
with Inflorescences single,
percurrent reticulation.
from
on branches and on older branches (then extending a structure
leafy (leafless)
axillary
mm
comprising the woody remnants of old inflorescences); short axillary shoot c. long,
1
mm
mm
mm
mmdiam.
5-7
diam. flower), 1-3 long, 2-2.5 (in fmit); pedicels long, c.
(in
1
mm
mm
mm
diam. the base, 2 diam. the apex (in flower), 8- 5 (20) long, 2-2.5
at at 1
.5
1
mm
mm
diam. the base, 3-4 diam. at the apex (in fruit); short axillary shoot sparsely
at
mm
lower broadly
covered with golden hairs long, pedicels glabrous; single bract
0.
1
mm,
1-2 by 1-2 acute, mostly caducous in fruit, rather densely covered with
triangular,
mm
golden hairs 0. long; upper bract inserted within basal half of pedicel, broadly trian-
1
mm,
1-2 by 1-2 glabrous; flower buds depressed ovoid; flowers green maturing
acute,
gular,
yellow in vivo, black in sicco, surfaces of sepals and petals glabrous, margins ciliate
to
mm
by
with golden hairs 0. 1-0.2 long; sepals basally connate, deltate, notreflexed, c. 3 c.
mm,
mm, 10-15 by 8-12
rounded, caducous, rarely persistent; outer petals ovate, inner
3
mm
mm;
androecium 6-7
12 by 6 receptacle depressed ovoid; diam.,
petals elliptic, c. c.
mm
mm
stamens long, connective appendage of inconsistent and irregular shape, 0.5
c. 1
Monocarps
number, and indument unknown. 10-22, green
wide, glabrous; carpels length,
maturing through red to black in vivo, black in sicco, ellipsoid, slightly asymmetrical, 2- 14
1
mm
mm
long, 7-8 diam., apicule ex-central, glabrous or sparsely covered with very short
mm
mm
golden hairs; stipes green maturing to red in vivo, 11-12 long, c. 1 .5 diam. increasing
mm
diam. when mature, glabrous or with sparse indument of very short golden
3
to
c.
mm
fruiting receptacle depressed ovoid, 5-10 in diam, glabrous. Seeds ellipsoid,
hairs;
mm
reddish-brown with small black surrounded by a slightly raised rim, 9- 3 long, 6-
pits
1
mm
unknown.
7 diam., raphe sunken, encircling seed longitudinally, rumination
Cremastosperma yamayakatense found only watershed
Distribution: in the
is
Maranon Amazonas
of the upper Rio in the department of in Peru.
m
Habitat and ecology Primary and secondary between 200 and 1000
forest
:
November
yamayakatense has been collected flower and January-
elevation. C. in in
specimens have been throughout year except December
March. Fruiting collected the
and April.
of other specimens examined.
Selection (21)
PERU: Amazonas, Bagua: Along roadside from Chiriaco to Puente
km NE
43 Feb 1996
Venezuela, (by road) of Chiriaco, 5
(fl),
Barbour, 4432 (MO, U, USM); District of Imaza, 22 Sep
P.J.
1997 Chavez 70 Aguarana de Yamayakat, 14
A., E. (U);
(fr),
1994 Diaz C; 6851 USM); Imaza,
Jul (fr), S., et al. (U,
community Yamayakat, Jan 1995 Hodges,
et 111
(fl), V.; al.
(HUT); Imaza, community Yamayakat, 6 Mar 1995
District
(fl,
Jaramillo, N. 584 (HUT, U, USM); District Imaza, community
fr),
Yamayakat, trail to Putuim, 22 Nov 2003 (fr), Pirie, M.D.; et al.
60 (HAO, USM); Yamayakat, margin Maranon 16
U, of river,
Oct 1995 Rodriguez 724 (HUT). Condorcanqui:
R., E.
(fr),
River Cenepa, Huampami, 7 Aug 1978 Ancuash,
vicinity of
(fr),
1324 East Cenepa, South Aintami Creek on
E. (U); of of ridge,
15 1974 1588 Quebrada Basusinuk,
Jul Berlin, B. (U);
(fr),
Huampami, 2 Nov 1972 Kayap, 33
tributary of R. (U); Valley
(fr),
km
Santiago, Quebrada Caterpiza, 2-3 from community
of river
Caterpiza., 12 Jan 1980 Tunqui, 573 (MO); 8 Sep 1994
S.
(fr),
Vasquez, 19055 (MO).
R.; et
(fr), al.
Notes.Cremastosperma yamayakatense resembles two other species of
Cremastosperma; C. gracilipes, which has been collected in the departments of Napo
and Pastaza in Ecuador, Loreto in Peru and in adjacent Colombia, and C. cenepense,
also described here, from the Cenepa region of Amazonas, Peru, with which distribution
its
The
therefore overlaps. most important differences between C. yamayakatense and C.
gracilipes are the flowers. C. gracilipes characterised by flower buds which open
in
is
during development and which bear indument on parts. In contrast, the flower buds of
all
C
yamayakatense bear virtually no indument and appear to remain closed throughout
when
development, the petals only opening slightly the flowers are mature. Additionally,
C
more
the flowers of C. gracilipes are borne on longer slender pedicels than those of
C
yamayakatense. C. yamayakatense differs from cenepense in the shape of the leaf
base (acute in C. yamayakatense, cordate to subcordate in C. cenepense) and the
length of the stipes (longer than the monocarps in C. yamayakatense, shorter than the
monocarps The
in C. cenepense). lack of flowering material of C. cenepense makes
further distinction currently impossible.
C
m
Flowering and fruiting specimens of yamayakatense of around were
1.5
tall
observed in the province of Bagua, though specimens collected both area and
in this
particularly those collected further north into the province of Condorcanqui, in the areas
m
of the Rivers Cenepa and Santiago, have been recorded as reaching heights of 6 - 8
m
and in one case 20 Differences between collections from these two regions have
tall.
been observed: The leaves of Condorcanqui specimens are generally larger and the
fruits
have a indument whereas Bagua
slight those of the collections are glabrous. In the absence
of from Condorcanqui assumed
material specimens do
floral these
is that represent the
it
same
due
species to the short pedicel, leaf base shape (which excludes the possibility of
their representing specimens of C. cenepense) and leaf venation.
12
Etymology
Cremastosperma yamayakatense named community
after the
is
:
of Yamayakat,
the type locaHty of
the species.
&
Cremastosperma -
cenepense
Zapata
3. Pirie spec. nov. Fig. 9
2,
Typus: Rojas, 269 AMAZ, HUT, MO, USM);
R.; et al. (holo, U, iso,
Amazonas,
Peru: Condorcanqui: Cenepa
Prov.
Ri'o region,
community Mamayaque, Aug
1997
11
(fr).
C
A
Lamina
basi cordata vel subcordata distinctum. yamayakatense
fructibus
A
indumento
monocarpiis
vestitis et stipitibus brevioribus
differt. C. gracilipes
pedicellis brevioribus
differt.
m
Tree, 10 young twigs and
c. tall; petioles sparsely (axillary buds densely) covered
mm
mm
mm
with appressed golden
hairs c. 0.1 long. Leaves: petioles 4-7 long, 1-2
diam.; lamina narrowly 12-22 by 4-8 cm
elliptic, (leaf index 2.7-3), chartaceous, grey-
yellow green above, brown
light or yellowish green below, glabrous on both
base
sides,
mm
cordate to subcordate, apex shortly acuminate (acumen 8- extreme
long), obtuse,
1 tip
mm
primary vein raised over entire leaf length, 1-1.5 wide widest
at point, glabrous,
mm
secondary
veins 7-12, intersecondary veins occasional, distance between from 2-5
at
mm
base
the to 15-25(35) closer to the apex, angles with primary vein from 90-80"
the
at
base 60-50'
to closer to the apex, forming distinct loops, smallest distance between
loops
mm,
and margin
2-5 more
tertiary veins or less percurrent. Inflorescence of
single
mm
mm
flowers,
axillary; short axillary shoot, 2 long, 2 diam.
c. c. (in fmit); pedicels 8
c.
mm mm
mm
long, 2 diam.
c. at the base, 2.5 diam. the apex
c. at (in fruit), short axillary
mm
shoot and
pedicels sparsely covered with appressed golden
hairs c. 0. long; 2 lower
1
bracts, caducous; upper midway
bract attached along
caducous;
pedicel, flowers not
Monocarps mm
observed. brown
8-10, blackish
in sicco, ellipsoid, asymmetric, 14-15
mm
9-1
long, diam., apicule
excentric, rather densely covered with appressed golden
1
mm
mm
mm
hairs long; 7-8
c. 0. stipes long, c. 1.5 diam., rather densely covered with
1
mm
mm
appressed golden
hairs long;
c. 0. fruiting receptacle depressed ovoid, 4-7
1
mm
diam., rather densely covered with appressed golden
hairs c. 0. 1 long. Seeds ellipsoid,
mm mm
brown
golden
shallowly wrinkled
(immature), 12 7
c. long, c. diam., raphe sunken,
encircling seed
longitudinally, rumination
unobserved.
Distribution: cenepense
C. has, to our knowledge, only been
collected
in the
Condorcanqui Province of Amazonas
in Peru, the area of Cenepa
in the River, a
tributary
of the River Maraiion.
Other
specimens examined:
PERU: Amazonas,
Prov. Condorcanqui: Rio Cenepa Que-
region,
brada Nahem, 15 1974 Kayap, r075(U); Rfo Cenepa
Jul R.
(fr),
community Mamayaque, Aug
region, 9 1997 Rojas,
R.; et
(fr),
255 MO,
(HUT, USM).
U,
al.
C
Notes cenepense yamayakatense
is similar to C. also described here, and
: ,
therefore also to C. gracilipes (see above). differs in the shape of the leaf base
It
(cordate or subcordate as opposed to acute in C. yamayakatense and C. gracilipes),
the indument on the fruits (rather dense as opposed to almost always absent in C.
C
yamayakatense), and
lengths of the pedicel (shorter than that of gracilipes) and
stipes (shorter than those of C. yamayakatense).
which
of
a
it
Acknowledgements
Two MSc
students from the University of Wageningen, the Netherlands; Marleen
MDP
Botermans and Robin van accompanied
Velzen,
in Peru: their assistance ensured
MB
RvV
the success of the expedition. and were supported through from
grants
the
Levenswetenschappen' MB, RvV
'Stichting (grant 805-40.201) and the 'Miquel Ponds';
MDP
and by from Mennega
grants the 'Alberta and was
Stichting' the expedition
further
,
financed by the Utrecht University branch of the National Herbarium of
the Netherlands.
The number
help of a of people in Pern during the field expedition gratefully acknowledged:
is
Museo
the staff of the de Historia Natural (Uni versidad Nacional Mayor de San Marcos,
Magda
Lima), Chanco
especially Estela, Oscar Tovar Serpa, Asuncion Cano
Echevarria,
Hamilton Beltran Santiago and Haydee Montoya
Terreros; the staff of the Jardin Botani-
co de Missouri, Oxapampa,
Peru, especially Rocio del Pilar Rojas Gonzales; Abundio
Alva
Sagastegui (Universidad Privada Antenor
Orrego, Trujillo); the staff of the Herbarium
Truxillense, Rodnguez
Trujillo, in particular Eric Rodriguez and Medina
F. Victor Ibanez;
and the people of the Comunidad Aguamna de Yamayakat, Imaza, Apu
especially Efrain
Wisum
Yagkug, Ricardo Apanu Nampin, and Julio Saan Kasen. The herbarium
curators
USM
MO,
HUT,
of and acknowledged Maps
are for the loan of collections. were produced
ESRI made New
using data available by the York Botanical Garden's Basemap
'Digital
of the Americas'. Paul M. Maas gave comment
valuable on
J. the descriptions and
Lubbert Th. Westra
Y. translated the Latin diagnoses.
Literature cited
Anonaceen-Gattungen Acta Horti
R.E. 1930. Revision der Arten einiger
Fries, I.
Berg. 1-128.
10:
Anonaceen-Gattungen. Acta Horti Berg.
1934. Revision der Arten einiger
,
12:203-207.
phytogeographical connections
Gentry, A. H. 1982. Neotropical diversity:
floristic
between Central and South America, Pleistocene climatic fluctuations, or an
Andean Orogeny? Ann. Missouri Bot. Gard. 69: 557-593.
accident of the
&
Myers, N.,C.G. Mittermaier, G. A. B. da Fonseca, Kent. 2000. Biodiversity
J.
hotspots for conservation priorities. Nature 403: 853-858.
W. Maas, van der
M.D., L.W. Chatrou, R. H. Erkens, T. Niet, B.
J.
Pirie, J. J.
&
Richardson. In Phylogeny reconstruction and molecular
Mols, E. Press.
J.
Annonaceae: of taxon sampling
dating four Neotropical genera of the effect in
in
new
on
age estimations Plant species-level systematics: perspectives pattern
in
P
and process (Bakker, FT., L. W. Chatrou, B. Gravendeel, and B. Pelser, eds.)
Regnum
Vienna, 149-174.
Vegetabile 142, Koeltz,
pp.
Fig Department of Amazonas
1: in Peru. Distributions of all three species are denoted by black
C
Fig of Maranon
Detail basin Amazonas.
2: in Distributions of bullatum Pirie denoted by white
C
&
yamayakatense
triangles, C. Pirie by black squares and cenepense Zapata by
Pirie
black
circles.