Forest Ecology and Management 302 (2013) 346–353

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Forest Ecology and Management

journal homepage: www.elsevier.com/locate/foreco

Mauritia flexuosa palm swamps: Composition, structure and implications

for conservation and management
Bryan A. Endress a,⇑, Christa M. Horn a, Michael P. Gilmore b
a
Division of Applied Plant Ecology, Institute for Conservation Research, San Diego Zoo Global, 15600 San Pasqual Valley Road, Escondido, CA 92027, USA
b
New Century College, George Mason University, 4400 University Drive, Fairfax, VA 22030, USA

a r t i c l e i n f o a b s t r a c t

Article history: Swamp forests dominated by the dioecious palm, Mauritia flexuosa, cover vast areas of the Amazon
Received 22 October 2012 Basin and are poorly studied despite their recognized ecological and economic importance. This knowl-
Received in revised form 19 February 2013 edge gap confounds current conservation and management efforts. In this study, we documented over-
Accepted 29 March 2013
story structure and composition of M. flexuosa palm swamps (aguajales) as part of a broader effort to
Available online 30 April 2013
understand their ecology and assist in developing best practices for multi-use management for the
Maijuna, an indigenous group in Loreto, Peru. In 12 aguajales, we sampled all trees > 5 cm DBH and
Keywords:
all palm individuals P0.5 m in height and grouped species based on their economic, subsistence,
Peruvian Amazon
Palm swamps
and cultural importance. Results indicated that aguajales are more diverse and structurally complex
Forest composition and structure than previous general descriptions suggest: though dicots were twice as abundant as palms
Forest management (p < 0.001), palms were taller and larger (p < 0.001) and thus still accounted for nearly half of stand
Mauritia flexuosa basal area. The most abundant species provide numerous timber and non-timber forest products, with
Euterpe precatoria over half of individuals having a Maijuna-recognized use. As a result of their complexity and usefulness,
aguajales are under more stress than previously thought, as most previous work has focused solely on
the destructive harvest of M. flexuosa. Aguajales are clearly ecologically, economically, and culturally
important ecosystems, and their conservation and management will require the prioritization and bal-
ancing of the numerous demands by both wildlife and people. To accomplish this, a better understand-
ing of their composition and ecological interactions is needed.
Ó 2013 Elsevier B.V. All rights reserved.

1. Introduction tion of aguajales particularly around the city of Iquitos (Vasquez

Swamp forests dominated by the large, dioecious palm Mauritia
flexuosa L.f. cover vast areas of the Amazon Basin and are both eco-
logically and economically important (Vasquez and Gentry, 1989;
Kahn, 1991; Delgado et al., 2007). Locally known as aguajales in
Peru, they cover approximately 5 million hectares (BIODAMAZ,
2004) and play several important ecological roles, especially in
terms of providing habitat and food resources to wildlife including
birds, ungulates, primates, and fish (Brightsmith, 2005; Bodmer,
1991; Beck, 2006). Additionally, aguajales store significant amounts
of carbon in their waterlogged soils (Vegas-Vilarrubia et al., 2010).
Aguajales are also important for rural and urban economies,
particularly in northeastern Peru, where millions of fruit from fe-
male M. flexuosa are harvested annually for raw consumption or
made into beverages, ice cream, and other products (Padoch,
1988; Peters et al., 1989). Harvest is generally destructive, as adult
females are felled to collect fruit resulting in widespread degrada-
⇑ Corresponding author. Tel.: +1 760 291 5486; fax: +1 760 291 5428.
E-mail addresses: BEndress@sandiegozoo.org (B.A. Endress),
sandiegozoo.org (C.M. Horn), mgilmor1@gmu.edu (M.P. Gilmore).
and Gentry, 1989; Delgado et al., 2007). Concerns about destruc-
tive harvest and associated ecological impacts have prompted gov-
ernment agencies, non-governmental organizations and rural
communities to promote and develop sustainable management
plans (e.g. Bejarano and Piana, 2002; Manzi and Coomes, 2009).
Management plans tend to focus exclusively on the sustainable
harvest of M. flexuosa fruit, despite the fact that rural communities
rely on aguajales to provide a wide range of commercial, subsis-
tence, and cultural goods and services (Gilmore et al., in press,
Kvist et al., 2001; Murrieta and Ruiz, 2011) and that a diverse array
of wildlife make extensive use of this habitat (Beck, 2006; Bowler
and Bodmer, 2011; Brightsmith, 2005). Consideration of these var-
ied aspects should be incorporated into conservation and manage-
ment plans; however, multiple-use tropical forest management
efforts, especially those involving non-timber forest products or
non-market goods and services, remain rare (Guariguata et al.,
2010).
To develop integrated, multiple-use aguajal conservation and
management approaches, a solid understanding of their forest
ecology is needed. However, the ecology, structure, and composi-
CHorn@
tion of aguajales is poorly understood, as past research has focused

0378-1127/$ - see front matter Ó 2013 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.foreco.2013.03.051
 B.A. Endress et al. / Forest Ecology and Management 302 (2013) 346–353
primarily on the ecology and exploitation of the dominant over-
story species, M. flexuosa (e.g. Holm et al., 2008; Horn et al.,
2012; Manzi and Coomes, 2009). Studies characterizing aguajal
plant communities are limited (e.g. Kahn, 1991) and empirical data
are lacking, as are evaluations of non-M. flexuosa aguajal forest re-
sources (but see Kahn, 1988 for assessment of palm resources).
Aguajales have been broadly classified into two types (BIOD-
AMAZ, 2004, Kahn, 1988; Kahn, 1991): floodplain aguajales, found
along the margins of rivers and streams, and upland aguajales,
formed in poorly drained depressions within upland forest that
are inundated only by rainfall. Both types are located on soils char-
acterized by partially decomposed organic matter under water-
logged, anaerobic conditions. Floodplain aguajales cover large
expanses of the Peruvian Amazon, while upland aguajales are
smaller but widely scattered throughout the region, embedded
within upland forest (BIODAMAZ, 2004, Kahn, 1991). Beyond these
basic descriptions, aguajal forest structure and composition re-
mains poorly documented.
The Maijuna, an indigenous group inhabiting several river ba-
sins in Loreto, Peru, have recently expressed interest in developing
management plans to better manage the use and extraction of
aguajal forest resources due to concerns about degradation from
the destructive harvest of M. flexuosa fruit (Horn et al., 2012). Cur-
rent aguaje stands show clear evidence of past over-exploitation
with low densities of adult female M. flexuosa and heavily male-
biased sex ratios (3.5 males:1 female; Horn et al., 2012). Low M.
flexuosa seedling abundances associated with low densities of adult
females suggest that harvest in some stands has affected seedling
recruitment rates (Horn et al., 2012).
In addition to aguaje, the Maijuna rely on aguajales for a num-
ber of commercial, subsistence, and cultural products, goods, and
services (Gilmore, 2005, Gilmore et al., in press). For example, over
60 plant species found in aguajales are used for construction mate-
rial, food, medicine, cultural ceremonies, and other purposes and
aguajales are important hunting areas for 20 species of mammals
and birds (Gilmore et al., in press). Several species of edible beetle
larvae are also harvested from the trunks of the palms M. flexuosa
and Oenocarpus bataua Mart. found in aguajales (Gilmore et al., in
press). Thus, aguajales represent an important and resource-abun-
dant area for the Maijuna. However, little is known about forest
composition or abundances of non-M. flexuosa forest resources,
making it difficult to develop appropriate management and conser-
vation strategies.
In this study, we documented overstory structure and composi-
tion of upland aguajales as part of a broader effort to understand
their ecology and assist in the development of multiple-use forest
management plans. Special attention was given to overstory palm
species, as palms have special significance and use not only for the
Maijuna (Gilmore, 2005), but also for rural communities through-
out Latin America (Peters et al., 1989; Brokamp et al., 2011). This
information will provide an ecological baseline to inform forest
management and complements previous work centered on the
ecology, harvest, and management of M. flexuosa (Horn et al.,
2012) and the ethnobiological significance of aguajal ecosystems
(Gilmore et al., in press). This study also represents the first pub-
lished dataset describing overstory structure and composition of
these widely distributed, ecologically and economically important
tropical forest ecosystems.
2. Methods
In 2010, twelve upland aguajal stands were sampled within
Maijuna traditional lands of the Yanayacu River basin located in
northeastern Peru (Fig. 1). The stands were selected from those
identified during a previous participatory mapping project
347
(Gilmore and Young, 2010). All stands were within a half-day of
travel (by foot/boat) from the Maijuna community of Nueva Vida.
Plots ranged from 0.25 to 5 km from each other. Felling of adult fe-
male M. flexuosa individuals to collect fruit has occurred in these
stands for many years, and current stands reflect this, with male-
biased sex ratios; the proportion of adult males in the canopy ran-
ged from 56% to 100% (Horn et al., 2012), significantly different
than un-harvested populations, which have 1:1 sex ratios (Kahn
and de Granville, 1992).
In each stand, a 0.1 ha plot consisting of three circular subplots,
each with a radius of 10.3 m, spaced 5 m apart was established. We
determined plot location by delineating the perimeter of the aguaj-
ales and then establishing the plot in a representative area at least
10 m from the aguajal edge. All woody stems and palms P5 cm
diameter at breast height (DBH) were identified, and their height
and DBH recorded (hereafter ‘‘overstory’’). Species were then
grouped based on their economic, subsistence, and/or cultural
importance to the Maijuna as well as the mode of harvest based
on Gilmore et al. (in press). Given the ethnobotanical importance
of palm species in the region, we expanded our sampling for over-
story palm species to also include individuals <5 cm DBH; for these
species (Astrocaryum chambira Burret, A. macrocalyx Burret, Attalea
maripa (Aubl.) Mart., Euterpe precatoria Mart., M. flexuosa, O. bataua,
O. mapora H. Karst. and Socratea exhorriza (Mart.) H. Wendl.) we
measured all individuals P0.5 m in height within the plots. Vou-
cher specimens were collected and deposited in the Herbarium
Amazonense (AMAZ), Universidad Nacional de la Amazonia Peru-
ana, Iquitos, Peru.
Overstory structural and compositional metrics were calculated
using standard methods and formulae. Additionally, for the six
most abundant palm species, population density and size class
structure based on height classes were calculated to provide addi-
tional insight on these economically and culturally important spe-
cies. Two palm species, M. flexuosa and E. precatoria, were
abundant enough to evaluate variation in abundance and size class
distributions among plots. In this paper, plot level variation of pop-
ulations of E. precatoria is reported. Results of M. flexuosa sex ratios,
population structure, and abundance are found elsewhere (see
Horn et al., 2012). Due to lower abundances of the other palm spe-
cies, plot level data were pooled to describe their overall popula-
tion abundance and structure.
3. Results
3.1. Aguajal community composition and structure
A total of 1124 individual trees were identified and measured.
Stem density averaged 937 stems/ha (±72 SE), with dicot (woody)
species being more than twice as abundant as palms (Fig. 2; t-test,
p < 0.001). Stand basal area averaged 27.8 m2/ha (±1.95 SE) across
the sampled stands; however, despite the greater density of woody
individuals, stand basal area was nearly equally divided between
palms and woody species (t-test, p = 0.466; Fig. 2). This was be-
cause palms, on average, were significantly larger (in terms of
dbh) than woody individuals (t-test; p < 0.0001), a trend that was
consistent across the 12 stands (Fig. 3). Additionally, the mean
height of palms was significantly greater than the height of woody
individuals (t-test, p < 0.0001; Fig. 3). Thus, the lower canopy lay-
ers of the aguajales were dominated by numerous, woody individ-
uals while the upper canopy (>20 m in height) was dominated by
palm species (Fig. 4).
A total of 138 tree species were identified from 36 plant families
(see Appendix I for complete list). Species richness averaged 31.1
species/plot (±2.1 SE). A high Simpson’s Diversity Index was calcu-
lated (0.904), representing a high probability that two randomly
348 B.A. Endress et al. / Forest Ecology and Management 302 (2013) 346–353

Fig. 1. Map of study area.

chosen individuals are different species, indicating a diverse over-
story. Overall, stand composition was characterized by a handful of
common species, most of which were palms, and numerous dicot
species that were infrequent and smaller in size (Table 1 and
Appendix I). For example, 54 species (39%) were represented by
just one individual and 112 species (81%) had fewer than 10 indi-
viduals recorded in all of the stands. Conversely, the 10 most abun-
dant species accounted for 55% of all stems sampled and 67.5% of
the total basal area recorded. Despite the high densities of woody
trees, relative importance values, as calculated from relative den-
sity, relative frequency and relative dominance, were low with
only Hevea guianensis var lutea (Spruce ex Benth.) Ducke and R.E.
Schultes), Virola pavonis (A. DC.) A. C. Sm., and Ruptiliocarpon cara-
colito Hammel and N. Zamora having importance values over 10.
Despite the low richness of palms (9 of the 138 species), three of
the top four most abundant species were palms: M. flexuosa, E.
precatoria, and S. exhorriza. M. flexuosa was by far the most abun-
dant species in the stands and accounted for over 40% of the basal
area.
As noted, palms were a major component of the overstory. Den-
sities and pooled population structures of the six most abundant
palms are reported in Table 2. M. flexuosa and E. precatoria were
 B.A. Endress et al. / Forest Ecology and Management 302 (2013) 346–353 349

Fig. 2. Box plot of palm and woody stem density and basal area across stands Fig. 3. Mean (±SE) height and diameter at breast height (dbh) of palm and woody
(N = 12). Points indicate mean density and basal area respectively. P-values are species in the twelve sampled stands.
based on t-tests.

dominant, with other palms having lower densities across all the
size classes. E. precatoria densities varied across plots, averaging
1125 individuals/ha (±232 SE) and ranging from 150 to 2340 indi-
viduals/ha. Population size class distributions of E. precatoria var-
ied considerably among the 12 sampled stands (Fig. 5) and were
heavily skewed to smaller size classes, with only eight individuals
(<1.0%) >10 m in height. Additionally a number of stands had low
abundances of the smaller size classes.

3.2. Aguajal forest resources

Aguajales provide a wide range of timber and non-timber forest

products (NTFPs), with 57 overstory species (41%) recognized and
used by the Maijuna. This accounted for nearly half (47.5%) of all
overstory stems measured (Appendix I). Species identified as use-
ful tended to be the most abundant components of the overstory,
with four of the five most abundant species having economic, cul-
tural, and/or subsistence importance (Table 1). These four species
(M. flexuosa, E. precatoria, S. exorrhiza and V. pavonis) accounted
for 33.1% of all recorded individuals and 52.9% of the basal area
in the stands.
Fig. 4. Mean density (±SE) of palm and woody stem density among height classes.

3.2.1. Timber majority of which (22 species) are used for subsistence and utili-
Twenty-five overstory species, accounting for 33% of stems and tarian uses (e.g. construction materials, firewood) and not com-
17.5% of basal area (4.84 m2/ha), provide timber resources, the vast mercially sold (Appendix I; see Gilmore et al. in press for details).
350 B.A. Endress et al. / Forest Ecology and Management 302 (2013) 346–353

Table 1
Absolute and relative density, frequency and basal area of the 20 most abundant trees. Importance values (IV) were calculated by summing relative density, relative frequency
and relative dominance. Information on current species use was generalized from Gilmore et al. (in press). Commercial harvest of timber from dicot trees (e.g. V. pavonis) is
currently banned due to Maijuna community rules, though timber can still be harvested for subsistence use. Harvest is non-destructive unless otherwise noted.

Species Family Density Freq. Basal area Rel. Rel. Rel. IV Commercial Subsistence or cultural use
(ind/ (m2/ha) den freq dom use
ha)
Mauritia flexuosa Arecaceae 127.5 1.00 11.7 13.6 3.3 41.9 58.9 Fruitb Fruitb, leaves, petioles, larvaea
Euterpe precatoria Arecaceae 83.3 0.92 0.7 8.9 3.1 2.6 14.5 Palm hearta, fruita Fruita, leavesb, trunka, roots
Hevea guianensis var. lutea Euphorbiaceae 55.0 0.58 1.6 5.8 1.9 5.9 13.7
Socratea exhorriza Arecaceae 49.2 0.67 0.5 5.3 2.2 1.9 9.3 Trunka Trunka, stilt roots
Virola pavonis Myristicaceae 49.2 0.75 1.8 5.3 2.5 6.5 14.2 Trunka Fruita, seedsa, trunka
Ruptiliocarpon caracolito Lepidobotryaceae 44.2 1.00 1.3 4.7 3.3 4.7 12.7
Macrolobium angustifolium Fabaceae 30.8 0.67 0.2 3.3 2.2 0.7 6.2
Euphorbiaceae sp. 1 Euphorbiaceae 29.2 0.58 0.9 3.1 1.9 3.5 8.6
Oxandra euneura Annonaceae 29.2 0.50 0.2 3.1 1.7 0.6 5.3 Trunka
Tabebuia obscura Bignoniaceae 22.5 0.42 0.2 2.4 1.4 0.7 4.5
Eriotheca sp. 1 Malvaceae 20.8 0.67 1.0 2.2 2.2 3.6 8.0
Carapa guianensis Meliaceae 18.3 0.25 0.5 1.9 0.8 1.8 4.6
Tabernaemontana sp. 1 Apocynaceae 17.5 0.42 0.1 1.9 1.4 0.5 3.8
Tapirira guianensis Anacardiaceae 15.0 0.50 0.4 1.6 1.7 1.5 4.8
Buchenavia seriocarpa Combretaceae 14.2 0.50 0.5 1.5 1.7 1.8 4.9 Trunka, bark
Didymocistus chrysadenius Euphorbiaceae 14.2 0.08 0.1 1.5 0.3 0.3 2.1
Attalea maripa Arecaceae 12.5 0.42 0.6 1.3 1.4 2.1 4.8 Fruitb, leavesb, seeds, spathe, larvae
Oenocarpus bataua Arecaceae 12.5 0.42 0.4 1.3 1.4 1.4 4.1 Fruitb Fruitb, leavesb, larvaea
Oenocarpus mapora Arecaceae 12.5 0.33 0.1 1.3 1.1 0.2 2.6 Fruitb, leavesb, Petioles, trunka
Pterocarpus amazonum Fabaceae 11.7 0.67 0.3 1.3 2.2 1.1 4.6 Buttress roots
a
Destructive harvest: individuals are killed during extraction.
b
Harvest can either be destructive or nondestructive, depending on harvesting technique (e.g. felling vs. climbing) and/or size of individual.

Table 2
Pooled abundance and density measurements by size classes for the six most abundant overstory palm species. Data from M. flexuosa are from Horn et al. (2012).

Palm Species
Mauritia flexuosa Euterpe precatoria Socratea exhorriza Oenocarpus bataua Oenocarpus maripa Attalea maripa
Height Class (m) Ind/ha % Ind/ha % Ind/ha % Ind/ha % Ind/ha % Ind/ha %
0.5- 0.99 525 24 344 31 169 50 43 43 12 13 43 48
1.0 - 2.99 1,135 52 538 48 93 28 27 27 48 52 27 29
3.0 - 5.99 323 15 156 14 32 9 14 14 22 23 11 12
6.0 - 9.99 68 3 53 5 13 4 7 7 11 12 6 6
10.0 - 19.99 50 2 33 3 28 8 10 10 1 1 4 5
20.0 - 29.99 76 3 2 0 1 0 0 0 0 0 0 0
>30 11 <1 0 0 0 0 0 0 0 0 0 0
Total 2,188 1,125 335 100 93 91

Of the four species with commercial value, V. pavonis and S. exhor-
rhiza were the most abundant. Harvest of these two species results
in the death of the trees, as neither species resprouts following har-
vest. While V. pavonis was relatively abundant (55 stems/ha), the
population was dominated by small individuals as 63% of individ-
uals had a DBH of <20 cm.
the fruit. Felling of M. flexuosa and O. bataua also provides opportu-
nities to cultivate several species of edible beetle larvae from the
decomposing trunks. However, felling of these species may also
limit other subsistence or cultural uses.
4. Discussion and conclusions

3.2.2. Non-timber forest products 4.1. Forest structure and composition

The Maijuna recognize 44 species that provide NTFPs. Uses vary,
though the majority (N = 26) provided edible fruit for either subsis-
tence or commercial use. Other products include resin, medicine,
caulk, and construction materials (Appendix I; Gilmore et al. in
press). Many of the abundant species, including the palms M. flexu-
osa, E. precatoria, and O. bataua are important for commercial mar-
kets. Depending on the species and product harvested, NTFP
harvest may result in the death of the harvested individual (Ta-
ble 1). Palm heart extraction from E. precatoria always results in
the death of these individuals, and harvest is concentrated on large,
reproductive adults of this species. Fruit harvest from the palms M.
flexuosa and O. bataua can be done either destructively by felling
the tree, or non-destructively by using climbing devices to access
This study represents the first quantitative assessment of up-
land aguajal forest structure and composition. While our inference
space is limited to a relatively small area as compared with the
wide distribution of aguajales across the Amazon, it provides
important insights and a baseline for future efforts.
Our reported overstory basal area and relative dominance of
palms are similar with Kahn’s (1988) estimates of three floodplain
aguajales in the Ucayali River basin. Results also indicate that
although M. flexuosa is the most dominant species in terms of den-
sity and basal area, stands are not mono-specific, and exhibit con-
siderable structural and compositional complexity, with a wide
range of species and families represented. Due to the lack of
 B.A. Endress et al. / Forest Ecology and Management 302 (2013) 346–353 351

Fig. 5. Population size class distributions of Euterpe precatoria from the twelve sampled stands.

published information on aguajal forest communities, it is unclear
as to the reason for the high abundance of woody species docu-
mented. The vast majority of woody trees were small, and the
low abundance of these species in the upper canopy suggests that
while the species are able to recruit and establish in aguajales, the
waterlogged, anaerobic soils may restrict growth and survival rates
for many of the species, limiting their ability to reach the mid and
upper layers of the canopy. Destructive overharvesting may ac-
count for the extremely low abundances of some larger woody spe-
cies such as V. pavonis, which was heavily logged for many years.
Furthermore it remains unclear if the long-term destructive har-
vest of female M. flexuosa has affected stand dynamics in such a
way as to promote the establishment and growth of woody species
within this palm dominated plant community.
Stand densities and basal areas were also similar to reports
from non-aguajal Amazon floodplain and swamp forests (e.g. Nebel
et al., 2001; Wittmann et al., 2010). However, while aguajales are
perceived to be dominated by just one palm, M. flexuosa, our find-
ings suggest that the density, basal area, and importance values of
palms as a group, not just M. flexuosa, were considerably greater in
upland aguajales as compared to many other Amazonian wetland
forest ecosystems (Nebel et al., 2001).
352 B.A. Endress et al. / Forest Ecology and Management 302 (2013) 346–353
Because of the broad distribution of aguajales throughout the
Amazon, and large variation in the hydrologic and geomorpholog-
ical factors involved in their formation, more work is needed to
better document the composition and structure of not only upland
aguajales, but also floodplain aguajales, which are different in that
they are inundated by seasonal flooding from rivers and streams.
Thus, extrapolating our findings to other regions or conditions
must be done with caution; clearly more research is needed to bet-
ter understand aguajal ecology and the factors influencing their
structure and composition.
4.2. Implications for conservation and management
Results highlight why Maijuna communities value these eco-
systems. As noted, the majority of the most abundant species ac-
counted for a large portion of forest basal area and provide a
myriad of commercial, subsistence, and cultural products. In addi-
tion to plant resources, aguajales are important areas for hunting,
as many hunted wildlife species visit aguajales to consume M.
flexuosa fruit (Gilmore, 2005, Gilmore et al., in press), and game
meat provides a major source of income for the community (Horn
et al., 2012). Aguajal plant species reported here are not only useful
to the Maijuna, but have been recognized as important for many
communities throughout the Amazon (Kvist et al., 2001; Murrieta
and Ruiz, 2011), highlighting the fact that aguajales are utilized for
much more than simply M. flexuosa fruit, and management ap-
proaches to these ecosystems must reflect their multi-use status.
Because of the high abundance of useful species, over-exploita-
tion and degradation of aguajales is a concern. Harvest methods to
obtain timber and NTFPs from aguajales have generally involved
killing individuals, and aguajales near the Maijuna communities
clearly show effects of destructive M. flexuosa harvest, with low
abundances of adult female individuals and skewed sex-ratios
(Horn et al., 2012). The absence of large individuals of V. pavonis
and E. precatoria may also be the result of over-exploitation, which
Maijuna community members indicated occurred at an intense le-
vel between 2000 and 2007 for V. pavonis and at moderate levels
from 1990 through the present for E. precatoria. Observed variation
in these species may be due to environmental heterogeneity among
the stands, but given the history of intense destructive harvest in
the region, observed patterns are suggestive of overharvesting.
Destructively harvested species such as E. precatoria, M. flexu-
osa, and V. pavonis, as well as other species used by the Maijuna
(e.g. Oenocarpus spp., Attalea spp., etc.) are widely distributed and
harvested by communities throughout the Peruvian Amazon.
Therefore, if these species are also common components of aguaj-
ales throughout the region, harvest pressure on these forest com-
munities is likely intense and widespread as extraction is not
simply restricted to M. flexuosa, but rather due to resource extrac-
tion from many of the most abundant overstory species.
Destructive harvest not only affects the harvested species, but
also may affect a number of ecological processes and interactions
or the availability of other forest resources. Lower abundances of
palm fruit due to reduced densities of M. flexuosa, E. precatoria,
and other palms (e.g. Astrocaryum spp., Oenocarpus spp.), may af-
fect the abundance of wildlife, such as tapir, peccaries, and prima-
tes, that visit aguajales to consume fruit of these species (Bowler
and Bodmer, 2011, Gilmore et al. in press). Notably, some of these
species are thought to influence aguajal dynamics through seed
predation and dispersal (Bodmer, 1991) leading to further disrup-
tion of ecological interactions. Additionally, given the importance
of palms to overstory structure and composition, it remains un-
clear if reduced palm abundances due to destructive harvesting af-
fects stand dynamics or is the cause of the relatively high densities
of woody species in the stands.
Recently, the Maijuna have taken steps to reduce over-exploita-
tion of aguajales. In 2007, commercial extraction of timber from
their traditional lands was banned, reducing pressure on V. pavonis
and other commercial timber species. Additionally, they have
worked to implement non-destructive harvest methods for collect-
ing fruit from M. flexuosa and O. bataua in an effort to increase com-
mercial production and to promote game species recovery, which
is an important part of their livelihood (Gilmore et al., in press,
Horn et al., 2012). Developing management plans for E. precatoria
harvest is more difficult, as destructive harvest is unavoidable
and this species is poorly suited to harvest (Zuidema and Boot,
2000). Given the low stocks of harvestable E. precatoria, their slow
growth rates, and the low price of palm heart, it may be that their
role in providing fruit for wildlife outweighs the limited income
gains through continued harvest. Efforts by the Maijuna have cer-
tainly begun to address aguajal conservation and management;
however, given the wide range of important resources and likeli-
hood of tradeoffs between different management goals (e.g. man-
aging for sustainable commercial fruit harvest vs. managing for
high abundances of game species), more must be done to strength-
en and optimize current efforts and develop management ap-
proaches that address a wide range of ecological, economic, and
cultural considerations.
4.3. Conclusion
Aguajales are complex systems that are heavily relied upon by
people as well as wildlife, and approaches to their conservation
and sustainable management should reflect the multiple nature
of the ecosystem and not simply be restricted to management of
M. flexuosa. Yet, our understanding of aguajal forest composition
and structure as well as the factors affecting forest dynamics is
lacking, confounding efforts. What is clear is that upland aguajales
are more diverse than previous general descriptions suggest and
are under more stresses than simply the threat of destructive M.
flexuosa harvest. Given the multiple uses of aguajales it is clearly
important to develop a process by which key resources and ecolog-
ical interactions are prioritized and balanced to support the con-
servation and management of these ecosystems.
Acknowledgements
We thank the Federación de Comunidades Nativas Maijuna (FEC-
ONAMAI) for their collaboration. S. Ríos Ochoa, V. Ríos Torres, D.
Ríos Vaca, L. Mozoline Mogica, E. Mogica Ríos, and A. Mozoline
Mogica provided assistance in conducting field research. E. Valder-
rama Sandoval and V. Vargas Paredes assisted in collecting data.
Research was conducted with the approval of FECONAMAI and
Nueva Vida. Botanical specimens were collected under Peruvian
permit N 0388-2010-AG-DGFFS-DGEFFS. C. Grández Ríos provided
assistance throughout the course of this project. Rainforest Conser-
vation Fund and Nature and Culture International provided in-
country logistical support.
Appendix A. Supplementary material
Supplementary data associated with this article can be found, in
the online version, at http://dx.doi.org/10.1016/j.foreco.2013.03.
051.
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