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Biodiversity and Conservation 3, 227-241 (1994)

Diversity and conservation status of Peruvian palms


FRANCIS KAHN" and FARANA MOUSSA
ORSTOM, CP-09747, 70001-970 Braslia (DF), Brazil

Received 24 May 1993; revised and accepted 3 September 1993 Indigenous palm species of Peru are listed with data on their distribution patterns, ecology, frequency. density in the ecosystems, and conservation status. Peruvian palm flora includes 140 native species in 34 genera with the following distribution patterns: strictly Andean (17), Andean and Subandean (3), strictly Subandean (19), Subandean and Amazonian (20), strictly Amazonian (7&), Amazonian and South peripheral ( 2 ) , South peripheral (1). About 43% of the species occur at very low or low frequency in the country and about 9% are insufficiently known in situ for their conservation status to be defined. There are no Extinct species. Sixteen of the 17 strictly Andean palms are threatened species; 3 of them are Endangered, while only 5 strictly Subandean, 3 Subandean-Amazonian, and 4 strictly Amazonian palms are in these categories.

Keywords: Palmae; Peru; distribution patterns; ecology; conservation status


Introduction

Macbride (1960) published the only palm flora for Peru. It is now out of date. Shortly after its appearance, Harold Moore organized an expedition throughout Andean and Amazonian Peru (Moore et al., 1960). Since then, several genera have been revised: Geonoma (Wessels Boer, 1968), Chelyocarpzls (Moore, 1972), Jessenia and Oenocarpus (Balick, 1986; Bernal et al., 1991), Hyospathe (Skov and Balslev, 1989), Dictyocaryzlm, Iriartea, Iriartella, and Socratea (Henderson, 1990), Ammandra and Phytelephas (Barfod, 1991), Astrocaryum (Kahn and Milln, 1992), Chamaedorea (Hodel, 1992), Aiphanes (Borchsenius and Bernal, in press); and two new genera were described, Itaya (Moore, 1972) and Aphandra (Barfod, 1991). Several taxonomic up-datings (Gentry, 1986; Kahn, 1990) and floristic inventories of palms in forest ecosystems (Kahn and Meja, 1990, 1991;'Kahn and Granville, 1992; Young 1992) were also published. In addition, countless specimens have been collected by R.B. Foster, A. Gentry, N. Jaramillo, F. Kahn, K. Meja, D.N. Smith, R. Vsquez, K.R. Young, and many other botanists in the last fifteen years. An analysis of a database including about 3000 herbarium specimens has provided information on species identification rates in each genus, species distribution patterns, and palm collecting intensity throughout Peru (Kahn et al., 1992). All specimens in each Subandean and Amazonian species have been listed according to the main river valley where they were collected, and never or poorly-collected areas were identified as a result (Moussa et al. , 1992).
"To whom correspondence should be addressed.
0960-3115 01994 Chapman & Hall

O. R.S.T.0A Fonds Documentaire


No?
Cote

O 8 SEP. 1994

qOLo?/2 : 6

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A list of indigenous palm species of Peru is presented with data on their di\tribution patterns. ecology, trequency and density in the ecosystem\. Their conservation status is defined from these parameters.
Materials and methods

.$pcit?s The list of indigenous palm species of Peru has been estnblished from idrntific*ation of herbarium specimens (AMAZ. BH. CUZ, K, MO. MOL. NY. P. LJS. LJSM- acronyms t h e checklist according to Holmgren tt cil., 1090, Tndex herharorium. 8th ed.). It follow~s of Peruvian palms prepared by L. Brako. A. Henderson and F. Kahn, and revised by J. Dransfield. in Brako and Zarucchis catalogue of flowering plants and gymnosperms of Peru (in press). Herbarium voiicher reference\ will he found in this catalogue and in Kahn and Moussa ( 19Y4).
List of
Pviti
pcdiri

Dimibiirioti p a t t t v x ~ of Feriii?rtri pibris

Five distribution patterns are propohed: ( I ) Strictly Andean species c)ccur at high elevation ( > 15V(lm) beyond the western limit of the Amazonian drainage which is marked by the occurrence of Dictyoc(iryit?i1mt~inl.ckirrtiiriri:Ctjro.xyloti spp. are typically Andem palms. they reach the western piedmont ot the Andes in northern Peru near the frontier with Ecuador: (2) Andean and Subandean \pecks: ( 3 ) drictly Subandean species occupy the eastern piedmont o f the Andes; (4) Amazonian specie4 are found in the plain (strictly Amazonian). some of them reach the piedmont valleys below 1000 m (Subandean-Amazonian) or extend beyond the southern limit of Amazonia ( Amazonian-South peripheral); ( 5 ) South peripheral species are found in b l adre de Dios; they dominate tree vegetation in savannas and gallery forests which extend to Beni. Bolivia. and muth of Rondhia. Brazil.
Pcriwiriii ecosystcs with pcrlrm

Ten ecosystems are con4idered: ( i ) Terra firme forests o n clay, usually well-drained soils with the highest palm diversity; (ii) dry white sands with low vegetation and very low palm diversity; (iii) waterlogged white sands with high palm diversity: (iv) seasonal swmip forests irregularly flooded hy rainfall with high density and medium diversity of palms: (v) permanently flooded swamp forests usually composed of very dense populations of hfrriiririrrj7e.i iioLsci.but a rather low diversity in palms: (vi) periodically flooded whitewater forests on alluvial soils (called restinga-forests in Peru. vh-zea-forests in Brazil) with medium palm diversity; (vii) fore5ts periodically flooded by blackwater with low palm diversity (called tahuamp~i-forestsin Peru. igapci-forests in Brazil); (viii) s;viinnas a n d (ix) gallery forests, both with low palm diversity: (s) mountain cloud forests above 1500 m. Some species occur in forests flooded by whitewater as well as in those floodecl by blackwater; they are considered riparian i11 the list. The ability of some torest palms to flourish in deforested areas is noted under open vegetation. Descriptions of these ecosystem\ in Peru are given by Marmillod ( 19x2). Encmucin (108.5). Knlliola c r d.(1987). Lamotte (1990). and Lopez and Freitas (1I9YO) for Amazonian lowlands. and by Weberbauer (1945), Ferreyra ( 1950). ONERN (1976). Young and Le6n (1988). and Young (1990) for eastern piedmont and the Andes.

Diversity and conservation of Peruvian palms

229

The ecosystem(s) given in the list is (are) that (those) where the species is commonly found. The attribution of only one ecosystem to a species does not exclude a certain ecological range within this species, which cannot be taken into account here. For instance, Astrocaryum javarense or Chelyocarpus repens form very dense stands on welldrained soils in terra firme forests; some individuals, however, may occur in the contiguous seasonal swamp forest. More information on species richness, density, vertical distribution, life forms, and ecology of palms in Subandean and Amazonian forest ecosystems are proposed by Kahn and Granville (1992). Data on Andean palm flora will be found in Young (1992).

Species ecology, frequency i n Peru and density in ecosystems The attribution of lhe above-defined ecosystem(s) to each species, as well as the frequency and density evaluation are mainly based on the field experience of the senior author who has been working with palms for 8 years in Peru. The species which have not been seen in the field are mainly those which are known only from the type or from a few old specimens (most often destroyed at B). As far as possible, complementary information was supplied from specimen labels, ecological and floristic studies (Gentry, 1985; Young, 1992), and forest inventories. Many of the latter, however, do not take palms into account, and when they do, the frequent absence of herbarium references make most identifications questionable. Conservation status The following IUCN conservation categories (Dransfield et al., 1988) are employed: Extinct, Endangered, Vulnerable, Rare, Indeterminate for threatened categories; Status unknown, Insufficiently known for unknown categories; and non-threatened categories. The conservation status of each species incorporates its distribution pattern because of the difference of human impact between regions, its frequency in the region, and its density in the ecosystem. As a general rule species which occur in very low frequency and are not known from other countries, or are known in very low frequency therein, are included in threatened categories: (i) as Endangered when there is evidence that current populations are mere remnants of former larger populations under strong human pressure; (ii) as Vulnerable when population density is low and the deforestation rate is still high; (iii) as Rare when population density is high because the species generally grow in open areas and can persist under strong human pressure; (iv) as Indeterminate when human pressure is low. Andean and Subandean species with low frequency and low density are also included in the threatened categories while Amazonian species with the same parameters are not. As a matter of fact, human pressure is very high in Andean and Subandean parts. Terrorism in the highlands made people migrate to the eastern Andes; mountain cloud forests have been intensively cut as a result. Many people were also attracted by coca cultivation on the piedmont where many areas have been deforested. Amazonian species, even with low frequency, may be considered not threatened as yet because of the lack of access roads and the consequently low density of human settlements in Amazonian lowlands. The unknown categories include species which have long been described and, since then, have not been found again, e.g., Catoblashu pubescens, Geononia congestissinia, G. dicranospadix or Wettinia weberbaueri.

Results

Ditwsity of Pci irviciii yalins A total of I40 native species in 31 genera has been listed (Table I ) . Five species which . EIiiei5 gziii~twzsis, are not indigenous palms of Peru are excluded: Bacfris gasipat~, Eirteipe olovctw. Oniocarp1rs htrcaba. rind Pliotwix cmitiririisis. Di&trihiitiotipiitttv I I Y Strictly Amazonian palm species represent 55.7"4 of the I40 species listed; 1'1.600 of them are strictly Suhandean species: 14.3'% of them are common to Amazonian lowlands ancl Suhandean peidmont. Andean specie? represent 12. l'h of the total: only three species are common to Andean and Subandean regions. Two Amazonian species also occur beyond the 5outhem limit of the hasin (Amazonian-South peripheral palmi,); there is only one South peripheral species (Table 2).
Ec)J ~ . ~ t t ~ i i 1 ~ All Andean and severiil Suhandean species occur in mountain cloud forests. A total of 70 Amazonian and Suhandean species grow in terra firme forests. and Ill of them are alscr found in other ecosystems. There are 16 nnd 27 5pecies in forests on periodically flooded alluvial wils and in seasonal swamp forests. respectively: six of the former and 17 (of the latter present a wider ecological range, however. The other ecosystems have a few palm species. but these n i q he very abundant (Kahn :ind Granville, 1997). All species occurring in forests which are picrdically flooded by blackwater, such as A,w";viiiii jtrrrtrri. Btrcrris tmirrrjli. B. ripiriu. arc also found in most inundated forest types (see riparian species in Table 1). The ecology of five species is still unknlown.
FrtTq i r tv i qv (it i d rici i sir? Thirty two specie4 occur at very low trequrncy. 29 at low frequency. On tht whole, 43.h",, o f the p l r n 5pccies are scarce. and 70.5'k of these are ulso found at low density. Frequency and den4ty are unknown for 11 specie\.

Di stribiitiori ptrttmi l u i d fr tiqiitwcy All strict11 Andean species occur at Lery low or low treyuency (Table 2 ) . Scarcc species o f the palm flora uf the Andes. the eastern reprewnt 9O.O";i. 45.7%. and 31 piedmont. and Am:imnian lowlands. respectively. The South peripheral specie<,,A stroc~zrv11iri liii(zii~ii. was collccted once in Peru. hut it is abundant in the adjacent Bolivian region.
Cori wrtwtioii itntris There are n o Extinct palm species in Peru. Twenty eight (20.0%) of the specie\ are, however. listed a s threatened: 9.3% are unknown: and 70.7% are not threatened. Sixteen (94.1 % ) strictly Andean species are within the threatened categories. c't'ro.t.vloii Irrriscctuiii C. i~~rri~cirlosiriri. und c'. ~ v ~ h ~ r b a i iire r t ~considered i Endangered 5pecies because only small populations remain in sites where mountain forests are being intensively destroyed. C'~ro.xyloticrispim. which is les5 scarce, is considered Vulnerable. ,4 iphiics spiccitri m d Eiitc.rpt, ~ i r i i i i i i o ~ itwo i. species described recently. and 1( 1 species

'-'I -> Table 1. Checklist of Peruvian palm species with their distribution patterns, ecology, frequency in the country, density in the ecosystem, and conservation status
*'
1.
< I

Species

Distribution pattern" SNAM SNAM A SNAM SNAM AM

Ecosystemb

Frequency in Density in country' ecosystem' L V L VL VL L VL L L L

Conservation statusd nt R V'

$.
a

?. m

Aiphanes aculeata Willdenow Aiphanes deltoidea Burret Aiphanes spicata Borchsenius & Berna1 Aiphaiies ulei (Dammer) Burret Aiphanes weberbaueri Burret Aphandra natalia (Balslev & Henderson) Barfod Astrocaryum caniosunz Kahn & Milln Astrocaryum chambira Burret Astrocaryum chonta Martius Astrocaryum gratuni Kahn & Milln Astrocaryum huaimi Martius Astrocaryum huicungo Dammerex Burret Astrocaryriin jauari Martius Astrocaryum javarense Trail ex Drude Astrocaryuni niacrocalyx Burret Astrocaryuin perangustatum Kahn & Milln Astrocaryrim scopatuin Kahn & Milln Attalea tessinaniiii Burret Bactris acanthocarpoides Barbosa Rodrigues

TF
TF MF TF TF/DWS TF/O

L L

R
nt ntf

H
H

SA

PFAS TF/PFAS/O PFAS PFAS/GF SAV SSFIO

VL

Re nt nt nt ntf

AM AM/SP AM/SP
SP SA

H M M
VL VL

H H H
M H

Re
nt nt nt Re Re
? nt

AM AM
AM SA AM AM AM

R TF
TF TF/O PFAS TF TF

H M

M/H H

L
VL VL
?

H
H
?

L/M

N
W
CI

Species

Distribution pattern"

Ecosystem"

Frequeiicq in Densitq in countryL ecos~strmc hl


1

Conser\ ation $tatusd nt


>

1 3

Bac-tris iicmtitliospar~illa Trail ex Drude Bucrris myicstifidiu Dammer Bucrris ~iriridit~a~~tw (Trail) Drude Bwtris b@d[i hlartius Biictrk chlorririciitliu Poeppig ex hlartius Bactris coiiciiiriii Martius Biicrris corossilln Karsten Bucrris jistis~ (frons Marti us Bactris lirrtm Martius Buc.tris Iiirriilis (Wallace) Burret Bactris Xillipii Burret Buc.tris iviwuju hlartius Bucais trioiiricolu Barbosa Rudrigues Biictris piriiiiga Trail BucrriA ripiriii hlartius BiictriA ,iiriplic~fr.otis hlartius Btictris sphnerocirrpu Trail Bactris iitilis (Oersted) Bentham & Hooker f. es Hemsleq Cutoblastits clriciri Cook &: Doyle Catoblnstw pithesceils (Karsten) Wendland Cvroxyloti crispmi Burret Ceroxylori lutisectiiiri Burret

Ah1
AM A hI Ah1 Ah4

TF
>

hl

R
PFAS
>

hl
hI
1

nt
H
>

nt
3

AM Ah1 Ah4 Am X ha
Ah1

R SSF TF TF TF
TF

hl \L
)

hl L
1

nt nt
3

hl hl

L hl-H

nt nt nt nt nt nt
Et

ARI Ah1

R SSF
TF R TF

L H H
L hl H
hl hl

L h H
L h4 L

Ilt

Ah1 SA
Ab
AM'!

TF

O
TF

L
hl

nt nt

nt'

A A

hlF MF

L
VL

L L
I .
8

v
E'
.

A'

-I

Ceroxylon verrimdosum Burret Ceroxylon weberbaueri Burret Chamaedorea angustisecta Burret Clzamaedoreafragrans (Ruiz & Pavn) Martius Clzamaedorea linearis (Ruiz & Pavbn) Martius Clzamaedorea pauciJlrora Martius Cliamaedorea pinnatifrons (Jacquin) Oersted Chelyocarpus repeizs Kahn & Meja Clielyocarpus ulei Dammer Desmoiicus leptospadix Martius Demoncia longifolius Martius Desmoncus mitis Martius Desmoncus orthacanthos Martius Desinoizcus polyacanthos Martius Desmoncus setosus Martius Desnzoncus vacivils Bailey Dictyocaryum lamarckiaizum (Martius) Wendland Dictyocaryum ptariense (Steyermark) Moore & Steyermark Elaeis oleifera (Kunth) Corts Euterpe catinga Wallace

A A SAIAM SA SA SAIAM SAIAM AM SAIAM AM AM AM AM AM AM AM A AM

MF
MF
TF TF

VL V L

Ee
E"
nt nt nt nt nt

L
L
L M L VL V L M L

L L
L L

TF
TF SSF

H H H

TF
SSF PFAS

Ie
I
nt

H L
L L

K
nt nt nt nt nt nt R

WO TF
PFAS/O/R TF O/R h4.F

H
M

L L
L L H M

H
L L

L
V L

TF SSF

AM AM

H H

nt nt

WWS

Species

Distribution pattern
A

Ecosystemh

Frequency in Densit> in countryc eco>ystemL

Conservation statusd

Eiittvpe liiiiiitiosm Hender\on. G, d 1eano & klesn Eiitrrpt. ~ I E ntoritr L hlartius Gtiotioina ucrriilis hIartius Gcoiiotritr irndicolu Damnier es Burret Georioiria irriirzdiiiucwi hlartius Ceoriotiin nspidiijoh Spruce Gcorioiiiu liticzdijrru (Poiteau) Kunth Ct~oi~otnn bartlcttii Burret Ceonorrio hurrletrii Danimerex Burret Grorloinu brotigiiinrtii hlartius Georiomu cuiriutitr Trail Geotioriiu coiigestissirrin Burret
Geoiioriirr cziiitwtu Wendland es Spruce Grorionici deciirreris Wendland et Burret Linden ey Wendland Gtmiortitr d e i w w (Poiteau) Kunth

LIF

\L
H H
VL

v
nt
Ilt

.4hl Ah1 A AM Ah1 ARI

SSF
SSFiPSF hlF

H H L
51

le

TF Li SS SSF

hl

nt

L
I

L
I

I<
ntg

Ah1 SA Ah1 SA Ah1


Ah1
A

TF SSF
SSFTF TF

VL hl RI
)

L H hl
I

K nt
Ilt

>

TF
TF hlF

L
hl
VL

L
L L
hl
>

nt nt

\.
nt
)

Ah1

TF
I

H
>

Georioriim t1icrtrriosputli.v Burret Cwiiormr frrrirgiiirrr Wendland es Spruce Ceotioiin grcicipiles Damnier es Burrer Groiionio grtrtitiitrijiigu Burret Georionui Iirlriiiiitlio~~ltiticr Burret

SA
SA SA AtSA A

TF TF %IF LIF

L
?

L
1

Ilt

>

hl

\.L

!il L

Ilt

\.

-.
Geoiioma iiiterrupta (Ruiz & Pavn) Martius Geonoma juruaiia Dammer Geonoina jussieirana Martius Georioma laxiflora Martius Geoiioina lehmaiinii Dammer ex Burret Geoiioma leptospadix Trail Geoiionia liiideniaiia Wendland Geoiioina macrostachys Martius Geoiioma inarggrajja Engel Geonoina maxima (Poiteau) Kunth Geonoma megalospatha Burret Geoiionia oligocloiia Trail Geonoma piscicauda Dammer Geoiioina poeppigiana Martius Geoiionia pycnostaclays Martius Geoiioma spixiana Martius Geoiionia tamandua Trail Geonoina tessmannii Burret Geoiionia trailii Burret Geoiioina triglocli in Burret Geoiioma trigona (Ruiz & Pavn) Gentry Geoiioma uiidata Klotzsch Geoiioma weberbaiceri Dammer ex Burret Hyospathe elegans Martius Hyospathe ulei Dammer Iriartea deltoidea Ruiz & Pavn Iriartella steriocarpa Burret Itaya anticoruin Moore Lepidocaryuin gracile Martius
SAIAM AM SA AM A AM A AM A AM A AM AM AM AM AM AM AM AM SA A A A AM SA SAIAM AM AM AM TF TF MFISSF PFAS MF TF MF SSFIPFAS MF TF M M M M VL

M M H M L M L M L M L L H

nt nt nt nt

V
nt

H
VL M V L M VL

V
nt

V
nt

MF TFISSF TF TF TF TF SSF TF DWS TF

V
nt nt nt nt nt nt

L H H H H
L ? M M VL V L VL

H H H
L
?

K
nt nt

MF
MF MF
TFISSF TF TFIMF TF PFAS TF

M M L

Ve V V
nt

L L

H
V L

M H
L M

H L H H H
M

V
nt nt

I
nt

Species

Distribution pattern Ah1 A hl

Ecosyste~n~ TF SSF
\ i S SSF PSF DWS

Frequency in Density in Conservation countrv ecos~cten~~ statusd hl L


hl hl

nt nt
Ilt

Ah1 SA!.Ahl Ah1 A hl


Ah1 SA~AR.1 SAfXhI Ah1 Ah1 Ah

L \H hI
hl
hl

H
VH

hl

ut Ilt nt Ilt nt nt
ntg

TF
TF SSFW\WO SSF,PSFIPFAS TF

hi-H
hl H H L H L

H
\L H L
H
hl

TF
TF

nt nt nt
nt

SA[Ahi
SAjAhl

TF
PFAS SSF hlF

H H
L
L L
hl

A/SA
AS4

hl

nt

hIF
TF

L
L

nt Ilt nt

SA
Ah1

PFAS
TF

M
hl

Ah1

Ilt

I'

Scheelea phalerata (Martius ex Sprengel) Burret Socratea exorrhiza (Martius) Wendland Socratea salazarii Moore Syagrus sancoila Karsten Syagncs smitliii (Moore) Glassman Welfia georgi Wendland ex Burret Wendlandiella gracilis Dammer Wettirtia augusta Poeppig & Endlicher Wettinia longipetala Gentry Wettinia niayriertsis Spruce Wettinia weberbaueri Burret

SAIAM AM SAIAM SAIAM AM SA AM SAIAM SA SA SA

PFAS

M M/H M

nt nt nte nt nt ntf nt nt

TF/S-PSF/PFAS VH TF TF TF MF H M

3.
B

? (D

L
VL

M L
L
M

2 2
n

9 <

$
%

TF

TF

H
VL M
?

M
L H
?

TF TF/MF
?

E s.

Ve
nt
?e

"Distribution patterns - A: Strictly Andean species; NSA: Andean-Subandean species; AM: Strictly Amazonian species; AWSP: Amazonian-South peripheral species; SA: Strictly Subandean species; SNAM: Subandean-Amazonian species; SP: South peripheral species. 'Ecosystems - DWS: Dry white sands; G F Gallery forests; MF: Mountain cloud forests; O: Open vegetation; PFAS: Forests on periodically flooded alluvial soils (vrzea-forest); PSF: Permanent swamp forests (Mauritia flexuosa swamps); R: Riparian species (including forests periodically flooded by blackwater); SAV: Savannas; SSF Seasonal swamp forests; TF: Terra firme forests; WWS: Waterlogeed white sands: ?: Parameter unknown. "Frequency and density - vL: very low; L: low; M: medium; H: high; vH: very high; ?: parameter unknown. dConservation status - E: Endangered; V: Vulnerable; R: Rare; I: Indeterminate; ?: Status unknown; K: Insufficiently known; nt: Not threatened. eEndemic species. 'These species, rarely collected in Peru, are not included in the threatened categories because they occur at higher frequencies in other countries. Aphandra natalia has been reported by Meja (1992) for Peru; it has also been collected near the Peruvian frontier in Ecuador (Borgtoft Pedersen, 1992), as well as in Acre, Brazil (Henderson 1126, NY). Ashocaiyuin ltuaiini has been collected in Madre de Dios, Peru (Albn 6936, USM). Bactris utilis is considered a synonym of B. gusipaes (Glassman, 1972). 'These species are not likely to occur in Peru - Geonoma baculifera and Oenocarpus minor are common palms in easted and central Amazonia, respectively. 'These species are treated under genus Azalea in Brako and Zarucchi (in press).
ur

s 3-

'Table 2. Diversity o f Peruvian palms, numbrr of species nith very low anci h v frequency, and numhrr of species in ILJCN categories in relation to distribution patterns. (Legend: cet' Tahle 1 )

Conservation status Numher Distribution patterns Strictly Anclrun Andt'an/Suh3ndran Strictl) Suhandean
SAlAhl
Ot

spccit.s

Frequency \L L
~

Threatened categories E V R I 3
13
(1

Not threatened nt
1

LTnknown
I)

Ti
(J

1s

7
I

o
h

Aniazonian Ah1
ARIISP

3 7

I 5 4 17

o
o
II 0

o o
2 3 0 2 o 2

o o
1

3 10
17

o
4

o
5

o
o o o
4

o
1

o
II

South prripheral Ttrtal

o
2v

o o o o

65 2
1
99

o
(1

o
3

32

15 7

o f G c o i r o i r i ( i which gron in mountain cloud forests at high elevation are also Vulnerrihle. Most of these last occur from Venezuela to Bolivia (Wessels Boer, 1968: Glxssman. 1973: Balslev and Moraes. 1989); the Vulnerable status may be severe. However. it combines ( i ) the high fragility of mountain forest ecosystem.,, (ii) the strong human prewire i n the\e regions. and (iii) the fact that such understorey species are unable to survive in cletcrrested areas. CAwrnitin It+iiii(iiiirii. luirleiiiciiici. and ( i . iii(irggr(ijf(i have heen cla\\ified a 4 Vulnerable for Colombia ( Berna1 . 1980). Dictyoc*cirvzoirImiiurt kiiiizziiiz is thc only Andean \?mies within the non-threatened categcrrie5. The $itxition is not so seriou\ for the strictly Subandean species. Only 5 of the 19 ified in the threatened categories: two of them. Hyo.sp(ithe rrlri and ltLtrrrrrri loirgipmiltr. are Vulnerahle. Four specie5 :ire within the unknown categories. and ten lire d e . Only 7.0",, of the \prcies which c)ccur in Amazonia art' considered threatened. However. cl \pecir\ which are not sulficiently knuwti yet to he clar\ified are likely to increase those threatened categorie.,.

Eir h i i s iii Endemic p a l m include 14 species (Tnhle 1): \ix ot them are 4trictly Andean specie4 (3 Endangered and 3 Vu1ner:tble): fi\e ~e strictly Subandenn species ( 1 Vulnerahle, 3 Rare specieh (not threatened): and tbro are and 1 Llnhnown): cine is ;I Suhliriclran-Arna~otii~n strictly Amazonian 4pecies (Indeterminate and Rare).
Discussion
Ptwii3iaii p d i i i

tiiiwvity

The high generic diver4ity of Perukiian palms is mainly due to the diversity of the Amazonian region. Thirty twcr of the 3S palm gcnera found in the Amazon basin and

Diversity and conservation of Peruvian palms

239

the Guianas are present in Peru (Kahn and Granville, 1992). Only Ceroxylon and Welfia are not found in Amazonia. The diversity at the specific level is certainly high. However, such data are clearly dependent on the degree of confidence of species identification. This can be considered high for those species belonging to the above-mentioned genera which have been revised recently, It is not the case of many species of some genera which do need re-assessment, such as Bactris which presents a low rate (55%) of identified vouchers (Kahn et al., 1992). The preceding authors also pointed out that many species of Bactris (about 30%, considering new taxonomic arrangements by A. Henderson, personal communication) and Geonoma (more than 50%) are represented by only one or a few specimens. If several species are found to have been infrequently collected, it cannot be excluded that others have been erroneously named. The material is not complete enough to check the identification with confidence in many cases. This suggests that the species list proposed here is longer than it will be when the Peruvian palm flora is updated. On the other hand, several new species have been described (Gentry, 1986; Kahn and Meja, 1985; Kahn, 1990; Henderson et al., 1991; Kahn and Milln, 1992; Borchsenius and Bernal, in press) and many species were reported new for Peru (Kahn and Meja, 1986, 1990, 1991) in the last years. For instance, Astrocaryum huaimi, Mauritia carana and Manicaria saccifera were recently collected. Moreover, many locations in Peruvian Amazonia have not been explored yet, as already stated. On the whole, erroneous names will be replaced by those of the species newly described or reported for the country, and Peruvian palm diversity is not likely to decrease significantly as a result.

Future of Peruvian palms Andean species are nearly all threatened. Current activities are for an increasing rate of deforestation. Chamaedorea and Geonoma are small palms in the understorey of mountain forests and will disappear with the forest. Ceroxylon are tall palms which persist in open areas, but they are commonly used as building material and consequently cut down. There are only a few roads crossing the Andes; the situation is still not catastrophic. When the economy of Peru recovers, new roads will be built to develop the country, this is an unescapable need for Peru, and that will be the end of many Andean species. The situation will be similar in Amazonia. There are few threatened species because of the low density of human settlements. When roads and colonization extend through Amazonian lowlands, vast areas will be deforested and many species will become scarcer and scarcer. The striking fact is that 43.6% of the palm flora occur at low frequency (53% including unknown species, which are likely to be still unknown because their frequency is very low). It is easy to imagine the example of Brazilian Rondnia being transposed to the Peruvian Amazonia resulting in a similar ecological catastrophe.
Acknowledgements This work was supported by the international agreement between ORSTOM, France and Museum of Natural History (UNMSM), Lima, Peru, with grant WWF-US 3322. We are particularly indebted to John Dransfield, A. Henderson, Dennis V. Johnson, and

2 4I

Kdrri and

Moltssil

Eduardo Llrrah for their valuable criticisms of t h e manuscript. We alto thank L. Brako. M. Hoff. K . Meja. . Milkin who have contrihuted to the project.
References
Balick. h1.J. ( 198fi) Systematics nnct economic botany of the Ot~rioc~rrr~itt,s-Jt~sst~ri~~i ( Palniae) complex. A f i z . Ecori. Bor. 3, 1-140. Balrlev. H. and hloraes. vi. ( I98Y) Siriopsis ILLY p~zlrnt~rtzs (IC Bolii,iiz. Aarhus: AAL1 Keports 20. Barfod. A.S. ( I Y Y l ) A monographic study of the subfoniily Phytelephantc~ideae (Arecaceac). Optwz Rot. 105, 1-73. Bernal. R . ( I Y N ) Endangerment of Colnmhi;in palms. Priticipt>s33. 1 13-28. Bernal. R.. G:ileano. Ci. and Henderson, A. (1991j Notes on Orrioctzr~iis. in the Colornhi;in Amazon. Rrittoriitz 43, 153-64. Borchsenius. F.K. and Bem:iI, R. (in press) A rtioriogrzp1i of' A i p l i r z i i c ~ s (Pdriiczrl. New York: Flora Nentropica blonograph. Borgtoft Pedersen, H. ( 1993) LJses :inJ management of rlphtziidr~ziitrfrrlitz (Pwlmaej in Ecuador. Hiill. Irist. fr. ct. m i d i i t * s 21. 741-53. Brako. L. and Zarucchi. J. (in press) Catalogue of the flowering plants and gymnosperms of Peru. Missouri Botanical Garden. Dransfield. J.. Johnson. D. and Synge, H. (1988) T h opa11ri.s qf flit, r i w world: A corist~riwrioit cm.ws. Camhridge: IUCN-WWF Plants Conservation Programme, Publication No. 2. Encarnacicin. F. ( 1985) Introducci6n a la flora y vegetacicin de la Amazonia peruana: cstatlio actual dc los estudios, medio naturul y ensayo de una clave de detcrminncicin de Ias formaciones vegetales en Ia llanura tiniaz6nica. C h i i ~ f o l k ~ 30, w 37-51. Ferreyra, R. ( 1Wl) Infomie botdnico de la exploracicin cientfica al valle del Huallaga. in Inforitic snhle c.1 Hiiallqcr pp. 177-31 6, Lima: OCHOA. Ministerio de Kelaciones Exteriores. Gentry. A . H. ( I 985) Algunos resultados preliminares de estudios bot:inicos en el Parque Nacional del hlanu. In Report hfizriir pp. 1-33. Lima: LJniversidad Nacional Agraria. Gentry. A.H. ( 1086) Notes o11 Peruvian palms. A m i . Afisvowi Bor. Gard. 73. 158-65. Glassman. S.F. ( 1977) A ra,i.yioii of B. E. Dtzhlgrtvi'.~ iridex ot',-liriericari pabris. Lehre: J Cramer. irriti tlic lrirrrteirior. New York: Flora Henderson. A. ( I Y W ) Armicme. frzrt I . Iiitr(~)ifiictiioii Neotropics Monograph 53. Henderscm. A . . Galeano. G. and Mesa. E. ( 1 Y Y 1 ) A new species of Eiitcvpt~(P;ilmae) from Peru. Rrittoriiiz 43, 178-80. Hodel, 1). R. ( 1993) C'liarritreiiarr~rr.Lawrence, Kan International Palni Society, Allen Press. Holmgren. P.K.. lieuken. W. and Schofield. E.K. ( 1990) Irifev Hrrhororiiini ftzrr 1. TIic hc~rhorio of rht. ~vorld. Edn. S. LJtrecht: Bohn. Schettema and Holkemn. Kahn. F. ( 1990) Las pnlnieras del Arboretum Jenaro Herrera (Provincia de Requeria, Departamento de Lorrto. Pen). C'ontribucitki al estudio de la f o r a y de la vegetaci6n de la Amazonia peruana. XVII. Crriiciollt~z45, 311-67. Kahn. F. and (irunville, J.J. de ( 1992) Paltiis iri j)resr c~cw.systerris of ;lrnn;oriiiz. Berlin: Springer Verlag. Ecological Series, Volume YS. Kahn. F. and hlejia. K. ( 1986) The Amcrican oil palm. Elwis o l t i i f w r , in Peruvian Amazcmia. Priticipes 30. 187. Kahn. F. and Mejia. K. (1988) A new species of C'~ic~1~~crcirrpii.s ( P a l m x Coryphoide..ie) from Peruvian Am;izonia. Principes 32. 69-72. Kahn. F. and hleju. K. (199(.I) Palni communities in wetland forest ecosystems of Peruvian Amazonia. For. ECCJI.hlanag. 33/34. 169-79. Kahn. F. rind Mejia. K. (l9Yl) The palni communities of two "terra tirme" forests in Peruvian Amrizonin. P r i t i c i p 35, 22-6.
(LI

Diversity and conservation of Peruvian palms

241

Kahn, F. and Milln, B. (1992) Astrocaryum (Palmae) in Amazonia. A preliminary treatment. Bull. Inst. fr. t. andines 21, 459-531. Kahn, F. and Moussa, F. (1994) Las palmeras del Per-colecciones, Patrones de distribucin geogrfica, Ecologia, Estatutos de conservacin, Nombres vernculos, Utilizaciones. Lima: Travaux de lInstitut franais dtudes andines, Volume 59. Kahn. F.. Henderson, A., Brako, L., Hoff, M. and Moussa, F. (1992) Datos preliminares a la actualizacin de la flora de Palmae del Pen: intensidad de herborizacin y riqueza de las colecciones. Bull. Inst. fr. t. andines 21, 549-63. Kalliola, R.. Salo, J. and Mkinen, Y. (1987) Regeneracin natural de selvas en la Amazonia peruana. I . Dinmica fluvial y sucesin ribereiia. Lima: UNMSM. Lamotte. S. (1990) Fuvial dynamics and succession in the lower Ucayali basin, Peruvian Amazonia. For. Ecol. Manag. 33/34. 141-56. Lpez Parodi, J. and Freitas, D. (1990) Geographical aspects of forested wetlands in the lower Ucayali, Peruvian Amazonia. For. Ecol. Manag. 33134, 157-68. Macbride, J.F. (1960) Flora of Peru. Part I, No. 2, Palmae. Field. Mus. Nat. Hist. Bot. 13, 321418. Marmillod, D. (1982) Methodik und Ergebnisse von Untersuchungen ber Zusammensetzung und Aufbau eines TerrassenWaldes in peruanischen Amazonien. Gttingen: Thesis Georg-August Univ. Meja. K. (1992) Las palmeras en los mercados de Iquitos. Bull. Inst. fr. t. andines 21, 755-69. Moore, H.E. Jr., (1972) Chelyocarpus and its Allies Cryosophila and [tuya (Palmae). Principes 16, 67-88. Moore, H.E. Jr., Salazar, A.C. and Smith, E.E. (1960) A reconnaissance survey ofpalms in eastern Peru. Lima: Agriculture division. United States operation mission to Peru, International cooperation administration (unpublished). Moussa. F., Kahn, F., Henderson, A., Brako. L. and Hoff, M. (1992) Las palmeras en los valles principales de la Amazonia peruana. Bull. Inst. fr. t. andines 21, 565-97. ONERN (1976) Mapa ecolgico del Per y gua explicativa. Lima: Oficina Nacional de Evaluacin de Recursos Naturales. Skov, F. and Balslev, H. (1989) A revision of Hyospathe (Palmae). Nord. J. Bot. 9, 189-202. Weberbauer, A. (1945) EI mundo vegetal de los Andes peruanos. Lima: Ministerio de Agricultura. Wessels Boer, J.G. (1968) The geonomoid palms. Verh. Kon. Ned. Akad. Wetensch. Afd. Natuurk., Tweede Reeks 58, 1-202. Young, K.R. (1990) Biogeography and ecology of a timberline forest in north-central Peru. Boulder: Thesis, Univ. of Colorado. Young, K.R. (1992) Floristic diversity on the eastern slopes of Peruvian Andes. Candollea 46, 125-43. Young, K.R. and Len, B. (1988) Vegetacin de la zona alta del Parque Nacional Abiseo, Per. Publ. Museo de Hist. Nat. Ser. B (Botnica) 34, 1-37.