Ecography 37: 902915, 2014

doi: 10.1111/ecog.00774
2014 The Authors. Ecography 2014 Nordic Society Oikos
Subject Editor: Jens-Christian Svenning. Accepted 21 February 2014

Distribution and abundance of tree species in swamp forests

of Amazonian Ecuador
Nigel C. A. Pitman, Juan Ernesto Guevara Andino, Milton Aulestia, Carlos E. Cern, David A. Neill,
Walter Palacios, Gonzalo Rivas-Torres, Miles R. Silman and John W. Terborgh
N. C. A. Pitman (ncp@duke.edu) and J. W. Terborgh, Center for Tropical Conservation, Nicholas School of the Environment, Duke Univ.,
PO Box 90318, Durham, NC 27708, USA. NCAP also at: Science and Education, The Field Museum, 1400 S. Lakeshore Dr., Chicago,
IL 60605-2496, USA. J. E. G. Andino, Dept of Integrative Biology, 1005 Valley Life Sciences Building no. 3140, Univ. of CaliforniaBerkeley, Berkeley, CA 94720-3140, USA. M. Aulestia and W. Palacios, Herbario Nacional del Ecuador, Casilla 17-21-1787, Avenida Ro
Coca E6-115, Quito, Ecuador. C. E. Cern, Escuela de Biologa, Univ. Central del Ecuador, Casilla 17-01-2177, Quito, Ecuador. D. A.
Neill, Univ. Estatal Amaznica, Puyo, Pastaza, Ecuador. G. Rivas-Torres, Wildlife Ecology and Conservation and Quantitative Spatial
Ecology, Evolution, and Environment Program QSE3-IGERT, Dept of Wildlife, Ecology, and Conservation, Univ. of Florida, 110 NewinsZiegler Hall, PO Box 110430, Gainesville, FL 32611-0430, USA. M. R. Silman, Dept of Biology, Wake Forest Univ., 1834 Wake Forest Rd.,
Winston-Salem, NC 27106, USA.

Research to date on Amazonian swamps has reinforced the impression that tree communities there are dominated
by a small, morphologically specialized subset of the regional ora capable of surviving physiologically challenging
conditions. In this paper, using data from a large-scale tree inventory in upland, oodplain, and mixed palm swamp
forests in Amazonian Ecuador, we report that tree communities growing on well-drained and saturated soils are more
similar than previously appreciated. While our data support the traditional view of Amazonian swamp forests as lowdiversity tree communities dominated by palms, they also reveal four patterns that have not been well documented
in the literature to date: 1) tree communities in these swamp forests are dominated by a phylogenetically diverse
oligarchy of 30 frequent and common species; 2) swamp specialists account for 10% of species and a minority
of stems; 3) most tree species recorded in swamps ( 80%) also occur in adjacent well-drained forest types; and
4) many tree species present in swamps are common in well-drained forests (e.g. upland oligarchs account for
34.1% of all swamp stems). These observations imply that, as in the temperate zone, the composition and structure
of Amazonian swamp vegetation are determined by a combination of local-scale environmental lters (e.g. plant
survival in permanently saturated soils) and landscape-scale patterns and processes (e.g. the composition and structure
of tree communities in adjacent non-swamp habitats, the dispersal of propagules from those habitats to swamps).
We conclude with suggestions for further research to quantify the relative contributions of these factors in structuring
tree communities in Amazonian swamps.

The scientic literature describing the tree communities of

Amazonian swamps is scant and largely reducible to four
points.
1) Tree communities in swamps are typically dominated
by one or more species of arboreal palms (Kahn and Meja
1990, Kahn 1991, Kahn and Granville 1992, and many others). This is most commonly the quintessential Amazonian
swamp palm Mauritia exuosa, but several other palms
and dicotyledonous tree species have also been observed as
dominants (Table 1).
2) Tree communities in swamps tend to be less diverse at
local scales than those in adjacent, well-drained forests
(Richards 1952, 1969, Ducke and Black 1953, Dumont
et al. 1990, Duivenvoorden 1995, 1996, Pitman et al.
1999, Duque et al. 2001, 2002, Romero-Saltos et al. 2001,
Ancaya 2002, Cern et al. 2003, Normand et al. 2006,
902

Cern and Reyes 2007, and many others; but see Cornejo
Valverde and Janovec 2006 for an exception). While
this low diversity is understood to be a consequence of
point 1, its root cause may be the small size and insular
character of swamp habitats (MacArthur and Wilson 1967,
ter Steege et al. 2001).
3) The oristic composition and structure of tree
communities in Amazonian swamps can be extremely
variable from site to site, both within and between regions,
and often more so than in adjacent well-drained habitats
(Duque et al. 2001, Romero-Saltos et al. 2001, Ancaya
2002). Tree communities in swamps also change over time
along a variety of successional pathways (Kalliola et al. 1991,
Roucoux et al. 2013). This spatial and temporal variability
in swamp forests is typically explained as a result of spatial
and temporal variation in ooding intensity, depth, water

Table 1. A list of tree taxa that have been reported to be the single most common species in at least one swamp forest inventory in western
Amazonia. Only inventories of trees 10 cm dbh were considered. Species followed by an asterisk have also been reported as common in
well-drained forest types in western Amazonia by a variety of authors, including Pitman et al. (2001), Duque et al. (2002), Dick et al. (2003),
Cornejo Valverde and Janovec (2006), and Rivas (2006).
Species
Palms
Mauritia exuosa

Euterpe precatoria
Iriartea deltoidea
Mauritiella armata
Mauritiella sp.
Oenocarpus bataua
Socratea exorrhiza
Non-palms
Ficus trigona
Lueheopsis hoehnei
Oxandra polyantha
Sacoglottis ceratocarpa
Symphonia globulifera
Tabebuia insignis
Sloanea sp.

Region
all regions

References

southern Peru
eastern Ecuador
eastern Ecuador
central Peru
northern Peru
northern Peru

Kahn and Meja (1990), Kalliola et al. (1991), Foster et al. (1994, 2000, 2001), Freitas
Alvarado (1996a, b), Grndez et al. (2001), Romero-Saltos et al. (2001), Ancaya
(2002), Cern et al. (2003), Fine et al. (2006), Vriesendorp et al. (2004, 2006, 2007,
2008), Cern and Reyes (2007), Honorio et al. (2008), Corrales Medina (2010),
Garca-Villacorta et al. (2010, 2011), Householder et al. (2012), this study, and many
others
Cornejo Valverde and Janovec (2006)
This study
This study
Foster et al. (2001)
Kalliola et al. (1991), Vriesendorp et al. (2004)
Vriesendorp et al. (2004), Garca-Villacorta et al. (2011)

southern Peru
southern Peru
southern Colombia
northern Peru
northern Peru
northern Peru
southern Peru

Ancaya (2002)
Gentry (1988), Foster et al. (1994)
Duque et al. (2001)
Corrales Medina (2010)
Pitman et al. (2003a)
Roucoux et al. (pers. comm.)
Ancaya (2002)

chemistry, and other abiotic factors (as in temperate zone

swamps; Kozlowski [1984], Casanova [2000]), and is
reected in various schemes for classifying Amazonian swamp
vegetation (Prance 1979, Klinge et al. 1990, Kalliola et al. 1991,
Junk and Piedade 2004, Ricaurte et al. 2012; for recent
reviews of classication systems see Junk et al. 2011, 2013).
4) Tree communities in Amazonian swamps often
feature species that possess adaptations to life in permanently
wet conditions. These include the ability of seedlings to
survive long periods underwater (De Vries 1997) and the
presence of pneumatophores (Granville 1974) or elaborate
stilt or aerial roots (see also Joly 1991, Armstrong et al.
1994, Jackson and Colmer 2005, Schngart et al. 2005, de
Oliveira and Joly 2010).
These four points have dominated the literature on
swamp tree communities for so long that the current paradigm of Amazonian swamps an environmentally harsh setting in which only a handful of highly specialized tree
taxa can prosper is now well established. According to this
view, Amazonian swamp forests are primarily assembled
from the regional species pool by the exclusionary action of
a strong environmental lter: the anoxic conditions in waterlogged soils, which permit the establishment of specialists
with morphological adaptations for surviving such conditions but kill all other species. Another set of lters related
to water chemistry, ooding intensity, peat depth, peat or
soil composition, microtopography, and other such factors
are understood to permit a given species to be dominant
in certain swamps or at certain times but not in others
(Junk 1982, 1989, Ancaya 2002).
While this paradigm is satisfying in some respects, it is
clearly undeveloped in others. One shortcoming is that it
gives precedence to processes operating at the local scale
(e.g. seed germination and seedling survival in swamps) over
processes and patterns at the landscape or regional scales

(e.g. seed input from adjacent forest types, the size and shape
of swamp forests, characteristics of the regional species
pool), and thus reects an outdated view of how ecological
communities are assembled (Ricklefs 2008 and references
therein).
The current paradigm of Amazonian swamps also
suers from the long-standing bias in studies of tropical vegetation towards highlighting dierences between samples
while playing down similarities (Pitman et al. 2001; see
also Gaston 2011). Since the tree communities in welldrained forests adjacent to swamps tend to be extremely
diverse and are rarely dominated to a large degree by a single
species, many researchers have been content to document
dierences between swamp and non-swamp forest types
while overlooking some striking parallels. For example,
the common perception of swamps as a very distinctive forest type sits poorly with the fact that at least ve of the
tree species reported to be top dominants in western
Amazonian swamps (Table 1) have also been reported as
common in well-drained forest types there.
In this study, inspired by Ricklefs (2008) call to understand how communities are assembled by examining the
distribution and abundance of the species they contain across
the broader landscape, we examined patterns of tree species
distribution, frequency, and abundance in swamp forests
and two adjacent well-drained forest types (oodplain and
upland) at a lowland Amazonian site in eastern Ecuador.
Our specic questions included: 1) are swamp forests at
our site dominated by a suite of frequent and locally
common tree species (an oligarchy) in the same way that
upland forests there are (Pitman et al. 2001, Maca
and Svenning 2005)? 2) How many of the tree species growing in swamp forests can be considered swamp specialists?
3) How many species in swamps have also been recorded
in well-drained forests, and how many are common there?
903

Answers to these questions allow us to partition the tree

communities growing in swamps into distributional guilds as
follows: x% of stems belong to species that specialize strictly
on swamps, y% to species that are common in all three forest
types, z% to species that prefer well-drained forest types
but occasionally spill over into swamps, and so on. In turn,
that partitioning should provide some new ways to
think about why certain species are common in swamps and
others are not, why the common species in swamps are so
dominant there, and why local-scale tree diversity is so much
lower in swamps than in well-drained forest types. As
Fine et al. (2010) have recently done for white sand forests, a
more general aim is to help place a forest type that is often
viewed as distinct and atypical into a broader context by documenting its connections with the surrounding landscape.

Study site and forest types

All swamp and oodplain inventories and all but two of the
upland inventories were carried out in the aseasonal lowland

Amazonian forests of Yasun National Park and the adjacent

Huaorani Ethnic Reserve in eastern Ecuador (hereafter
referred to collectively as Yasun; Fig. 1). The remaining
two upland plots were established in similar forest 100 km
west of Yasun (see map in Pitman et al. 2001). These sites
and their ora are among the best studied in the Amazon
(Herrera-MacBryde and Neill 1997, Pitman 2000, Bass
et al. 2010, Pitman et al. 2011).
According to a new ocial map of Ecuadors vegetation
(Mapa de Vegetacin del Ecuador 2012), the study area is
dominated by upland forests ( 79% of the total), with
smaller areas, mostly along rivers, covered by swamp forests
( 19%) and oodplain forests ( 2%; Fig. 1). Swamp forests
in our study were dened as tree communities growing
on peaty substrate that was saturated at the time of eld
work and that appeared to remain so year-round. These substrates were composed mostly of decaying plant matter
(a large portion of which appeared to be leaf fragments),
under which mineral soil was typically only encountered
after digging 20 cm (and often not encountered at all).
Where mineral soil was found in swamps, laboratory analyses

Figure 1. The location of the study plots in western Amazonia. (A) The location of Ecuador in South America. (B) The location of
Yasun National Park in Ecuador. (C) The northwestern portion of Yasun National Park (black) and the eastern portion of the Huaorani
Ethnic Reserve (labeled). (D) The locations of 13 of the upland tree plots (U) and all of the swamp (S) and oodplain (F) tree plots
discussed in the paper. Numerals after a symbol indicate that multiple hectares were inventoried in the same vicinity. Forest type maps are
from a recent nationwide classication of forest types (Mapa de Vegetacin del Ecuador 2012). Swamp forest shown here includes
both monodominant Mauritia exuosa swamps and mixed swamps.

904

indicated that it primarily consisted of clay and silt, as is the

case for most soils in Yasun (% clay % silt % sand;
Pitman unpubl.).
Thick, peaty substrates were never seen in upland forests
(which we dened as forests that were never ooded) or
oodplain forests (forests ooded for short periods during
high-water events and otherwise well-drained). All swamp
plots held some standing water during eld work, and
shared an additional landscape feature which we did not
observe in other forest types in Yasun: sturdy hummocks of
roots, organic matter, and mineral soil at the bases of many
trees, standing as much as 50 cm higher than the surrounding terrain and giving the impression of miniature islands
surrounded by peat- or water-lled depressions. Even these
hummocks, however, were well under the maximum water
levels in swamps. We did not measure substrate moisture
levels during the inventories, and do not have data on the
origin, chemistry, seasonality, or hydrology of the waters
that ood the swamps we sampled.
All swamp plots were 1 km from rivers and their borders at least 50 m from swamp edges (i.e. from the nearest
non-swamp forest type). We did not consciously select certain kinds of swamps, but sampled what we found growing
on saturated soils within the vicinity of the 110-km pipeline
road that traverses the northwestern corner of Yasun. The
ocial map of Ecuadorean vegetation (Mapa de Vegetacin
del Ecuador 2012) distinguishes between two types of
swamp vegetation in Yasun National Park: one in which
nearly all trees are M. exuosa (presumably because ooding
there is severe enough to kill all other species; 66 418 ha)
and another in which M. exuosa coexists with other tree
species (presumably because ooding is not severe enough
to lter them out; 127 710 ha). All of our plots correspond
to the latter category. According to other systems for classifying swamp vegetation, the swamp forests we studied in
Yasun appear to correspond to palm swamps and forest
swamps sensu Kalliola et al. (1991), swamp forests
sensu Herrera-MacBryde and Neill (1997), lowland ooded
palm forest sensu Palacios et al. (1999), and forested
swampy wetlands (mixed forests) sensu Junk et al. (2013).
Because our study was carried out in a region where
neither blackwater rivers nor open, herb-dominated wetlands are common, our plots sample a relatively narrow
portion of the full range of Amazonian swamp vegetation
(Lpez Parodi and Freitas 1990, Kalliola et al. 1991). For
example, nutrient-poor swamps with dense, treeletdominated vegetation that have been described around
Iquitos, Peru (Encarnacin 1985, Garca-Villacorta et al.
2003, 2011), do not appear to occur in Yasun.

Methods
Field work
Field work was carried out at four sites in swamps
(19981999; Supplementary material Appendix 1), ve sites
in oodplains (19971999), and 15 sites in the uplands
(19871999; Fig. 1). At each site we carried out a standard
1-ha tree inventory of all free-standing trees 10 cm dbh,
with the exception of one swamp site (Tiputini 4) where we

established two adjacent 1-ha plots. All swamp and oodplain plots measured 100 100 m, but some upland plots
were irregularly shaped (Pitman et al. 2001).
In all plots trunk circumference at 1.3 m was measured
and stems permanently marked with numbered aluminum
tags. (In the Dicaro 3 plot only 200 of the 378 Mauritiella
armata stems were measured for circumference, because
their densely spiny trunks complicated measurements. The
basal area of that plot was estimated by assuming that all
unmeasured individuals had the mean circumference value
of the 200 individuals we did measure.) Circumferences
were converted to diameters by assuming that stem crosssections were round.
Trees that could not be identied in the eld were
collected and compared to mounted specimens in the
QCNE and QCA herbaria in Quito, Ecuador. This was
especially eective because the QCNE herbaria holds several thousand specialist-identied tree specimens collected
in Yasun. When a specimen could not be matched to
herbarium material, it was assigned a temporary morphospecies name and treated in analyses as though it were a
named species. A full set of vouchers from the tree plots
was mounted and deposited at QCNE and a subset deposited at MO. All swamp specimens were entered into
the TROPICOS database, under NP collection numbers
34663550 (Tiputini 4A plot), 39514052 (Tiputini 4B),
44564483 (Dicamunpare 1), 44844526 (Dicaro 3),
and 45284589 (Dicamunpare 2).
The upland plots described here represent the same
dataset described in Pitman et al. (2001, 2002), with the
exception of some taxonomic updates. To assign species to
plant families, we followed Angiosperm Plant Group conventions as of May 2010 (APG II 2003).
Analyses
To explore abundance patterns in the three forest types, we
asked 1) if an oligarchy of common and frequent species
dominated swamp forests, 2) how many species could
be considered swamp specialists, and 3) how many species
recorded in swamps were also recorded in well-drained forest
types (oodplains and uplands), and how many qualied
as oligarchs in those forest types.
Following Pitman et al. (2001), we dened upland oligarchs as species recorded in a majority of upland plots and
at an overall density of 1 individual ha1 in the uplands.
We used the same criteria to identify swamp oligarchs and
oodplain oligarchs. For example, swamp oligarchs were
species that were recorded in a majority of swamp plots and
at an overall density of 1 individual ha1 in swamp plots.
Species that qualied independently as oligarchs in all three
forest types are referred to in this paper as super-oligarchs.
To identify swamp specialists in this study we used two
arbitrary thresholds: one for strict specialists and one for
functional specialists. Strict swamp specialists were species
that had an overall density in swamps of 1 individual
ha1 and were never recorded in the other forest types.
Functional swamp specialists were species that had an overall
density in swamps of 1 individual ha1, for which 80%
of all individuals were recorded in swamps, and that did not
905

qualify as an oligarch in a well-drained forest type. Note that

because just 19.2% of all the stems we sampled in the plot
network were in swamps, a species with 80% of its individuals in swamps would have 94% of its individuals in
swamps if the same number of stems had been sampled
in swamps as in non-swamp forest types.
To check how the results of these arbitrary thresholds
compared with those of a commonly used method for identifying the dominant species in a given habitat, we performed
an indicator species analysis (ISA) with the full 25-plot dataset to identify species associated with swamps (Dufrne and
Legendre 1997). While the ISA provides some additional
context for the results of the arbitrary thresholds, we gave precedence to the latter due to their simplicity and transparency.
To summarize the results of the above analyses we quantied the proportion of all stems and of all species in swamps
that qualied for the three categories of distribution and
abundance described above: strict swamp specialists, functional swamp specialists, and super-oligarchs. Species that
did not fall into any of these three categories were classied
into one of three additional categories: too rare (species represented by 4 stems in the entire dataset and thus considered too rare for their distribution to be characterized),
spillover (dened arbitrarily as species that were represented
in the entire dataset by 5 stems, 13% of which were in
swamps), and other (for all other species). The 13% value
in the spillover denition comes from a binomial probability
calculation based on the fact that swamp stems account for
19% of the entire dataset; that calculation shows that species
with 13% of their stems in swamps are signicantly rarer
in swamps than expected (p 0.04).

Results
Structural and ecological attributes of the 25 tree plots are
summarized in Table 2, and additional details of the swamp

plots are given in Supplementary material Appendix 1. With

the exception of 34 trees that we were unable to identify, the
2983 swamp stems were sorted to 243 species-level taxa
(204 species and 39 morphospecies). Most morphospecies
were rare, and together they accounted for 5.6% of swamp
stems. Forty species recorded in swamps (16.5% of
all swamp species) were recorded in no other forest type,
and these accounted for 31.7% of all swamp stems. Thirty
of the species that were only recorded in swamps occurred
in a single plot there, and 17 of those were represented by
a single tree.
Swamp oligarchs
Tree communities in swamps were dominated by a strong
oligarchy. Thirty of the 243 species recorded in swamps
qualied as oligarchs there, and these accounted for
59.0% of stems and 12.3% of species in swamps. Within
individual swamp plots, swamp oligarchs accounted for
between 38.6 and 86.3% of all stems (average 61.0%).
The lowest value is from the Dicaro 3 plot, where nearly half
of stems belonged to a non-oligarch palm (Mauritiella
armata) which we recorded nowhere else. The most common
swamp oligarchs were the palms Iriartea deltoidea (ranking
no. 1 in abundance in two plots) and Mauritia exuosa
(ranking no. 1 in two plots; Table 3). Despite the clear
dominance of palms, the 30 swamp oligarchs were a
taxonomically diverse group, including 28 genera in 16
families (Supplementary material Appendix 1).
Although a strong oligarchy was observed in swamps,
this should not be misinterpreted to mean that community
composition and structure were uniform between swamp
plots. Both composition and structure were more variable
between swamp plots than between plots in other forest
types, as measured by the Sorensons and BrayCurtis indices
(Table 2).

Table 2. Structural and ecological attributes of the 25 1-ha tree plots studied at Yasun, Ecuador.
Attribute
No. ha sampled
No. stems sampled
No. species
No. families
Mean stems ha1
Mean species ha1
Mean basal area ha1 (m2)
% stems that belong to single most common species
% stems in 5 most common species
Abundance rank of Iriartea deltoidea
No. forest type oligarchs
% spp. that are forest type oligarchs
% stems that are forest type oligarchs
No. spp. recorded in every plot of the given forest type
No. singletons
% spp. that are singletons
No. spp. recorded in just one plot
No. palm species
No. palm species that are forest type oligarchs
% stems that are palms
Mean Sorensons index between plots within forest type
Mean BrayCurtis index between plots within forest type

906

Swamps

Floodplain

Uplands

5
2983
243
39
597
74
25.7
12.7
52.1
3rd
30
12.3
59.0
2
109
44.9
162
10
6
57.4
0.244
0.239

5
2730
511
53
546
189
27.9
6.6
14.0
1st
88
17.2
55.2
13
186
36.4
257
11
7
12.0
0.348
0.297

15
9809
986
70
654
239
30.2
7.8
14.7
1st
150
15.2
63.0
6
128
13.0
343
9
6
9.9
0.376
0.332

Table 3. The ten most abundant tree species in ve 1-ha inventories in swamp forests in Yasun, Ecuador.
Tiputini 4A

ind.

Tiputini 4B

ind.

Iriartea deltoidea**
Mauritia exuosa** (S)
Attalea butyracea** (S)
Euterpe precatoria**
Sloanea bark (NP3478) (S)
Socratea exorrhiza**
Cespedesia spathulata (S)
Macrolobium angustifolium (S)
Sterculia apeibophylla
Astrocaryum murumuru var. urostachys

216
65
54
48
24
16
14
9
8
7

Iriartea deltoidea**
Attalea butyracea** (S)
Mauritia exuosa** (S)
Euterpe precatoria**
Isertia rosea (S)
Astrocaryum murumuru var. urostachys
Socratea exorrhiza**
Sterculia apeibophylla
Apeiba membranacea
Pterocarpus amazonum/rohrii cf.

122
92
56
53
24
19
16
12
9
7

Dicamumpare 1

ind.

Dicaro 3

ind.

Mauritia exuosa** (S)

Euterpe precatoria**
Macrolobium angustifolium (S)
Zygia glomerata cf. (S)
Buchenavia amazonia (S)
Hieronyma alchorneoides var. estipulosa
Hydrochorea corymbosa
Cecropia engleriana
Virola peruviana cf.
Euphorbiaceae nondescript (NP4465) (S)

170
108
44
44
11
11
10
10
10
7

Dicamumpare 2

ind.

Mauritia exuosa** (S)

Triplaris weigeltiana
Casearia uleana
Zygia coccinea var. oriunda
Euterpe precatoria**
Hieronyma alchorneoides var. estipulosa
Cecropia engleriana
Pterocarpus amazonum/rohrii cf.
Virola pavonis
Duguetia spixiana

68
36
28
27
23
19
18
17
12
12

Mauritiella armata (S)

Euterpe precatoria**
Oenocarpus bataua
Macrolobium angustifolium (S)
Sloanea bark (NP3478) (S)
Hieronyma alchorneoides var. estipulosa
Virola surinamensis
Lauraceae palermo (NP4484) (S)
Socratea exorrhiza**
Sterculia frondosa

378
95
71
64
18
13
11
11
10
10

Key
boldface top ten in multiple plots
palm
(S) swamp specialist

Only two species occurred in all ve swamp plots: Euterpe

precatoria (Arecaceae) and Macrolobium angustifolium
(Fabaceae). Most species recorded in swamps (66.7% of the
total) were recorded in just one swamp plot, and most of
these occurred as a single individual (Table 2). Nearly half
of all species recorded in swamps (44.9%) were singletons.
Swamp specialists
Twelve tree species qualied as strict swamp specialists,
and these accounted for 4.9% of swamp species and
30.0% of swamp stems. Nine other species qualied as functional specialists. Together, these 21 species accounted for
8.6% of swamp species and 43.6% of swamp stems (Fig. 2).
This small guild of swamp specialists was dominated by
two palm species: Mauritiella armata (12.7% of all swamp
stems) and Mauritia exuosa (12.0%; Fig. 3).
Indicator species analysis identied 11 species with a
positive, statistically signicant association with swamp
forests, and was thus more stringent in identifying swamp
specialists than our arbitrary method. These 11 species
accounted for 34.6% of all swamp stems. Eight of the
11 swamp indicators are among the 21 species we dened
as specialists (ve strict and three functional specialists;
Table 4). The other three swamp indicators (Euterpe
precatoria, Hieronyma alchorneoides, Symphonia globulifera)

Figure 2. The proportions of all species and all stems in ve

1-ha swamp plots in Yasun, Ecuador, that belong to four dierent
distributional guilds, or that were too rare (too rare) or too
eclectic (other) to classify. See the text for guild name denitions.

907

Figure 3. A diagram illustrating distributional patterns of tree species across three forest types in Yasun, Ecuador. (A) The proportions of
species recorded in each forest type in each of four distributional patterns. For example, the left-hand square represents all species
in swamps. The rectangular portion labeled also in oodplains and uplands indicates that slightly more than half of all species recorded
in swamps were also recorded in both oodplain and upland forests. (B) A schematic cross-section of the Yasun landscape illustrating
one commonly observed arrangement of the three forest types there. The gray area is peat and the stippled area is a river. (C) The proportions of all stems recorded in each forest type that belong to species in each of four distributional patterns. Within each forest type the
top four rectangles correspond to the same sequence of distributional patterns shown in that forest type in (A). The fth, bottommost rectangle represents unidentied stems. In some categories the most common species is indicated within a rectangle whose size is
proportional to that species abundance.

were too common in well-drained forests to qualify as

specialists under our arbitrary thresholds.
Sixty species in swamps (24.7% of all species) qualied as
spillover taxa, and these accounted for 3.4% of trees in
swamps. Sixty-three species (25.9%) were too rare for
their distributional patterns to be classied, and 91 (37.4%)
were distributed between swamps and other forest types in a
way that placed them in the category other. These two
categories accounted for 3.0 and 21.4% of swamp trees,
respectively (Fig. 2).
Of the 30 swamp oligarchs and 21 swamp specialists,
nine species qualied as both: ve strict specialists and four
functional specialists.

Species distributions across forest types

The great majority of species recorded in swamps (83.5%)
were also recorded in at least one well-drained forest type,
and these species accounted for 68.3% of all swamp stems
908

(Fig. 3). Most species recorded in swamps (53.1%) were also

recorded in both well-drained forest types, and these
accounted for 50.4% of all swamp stems.
A large proportion of species recorded in swamps were
common in well-drained habitats. For example, 30.8% of
all species recorded in swamps and 53.3% of species that
qualied as oligarchs in swamps also qualied as oligarchs in
one or both well-drained forest types. The 51 upland
oligarchs recorded in swamps accounted for 34.1% of all
stems there; comparable numbers for oodplain oligarchs
are 50 and 34.2%. Together, 75 species that qualied as oligarchs in at least one well-drained forest type were recorded
in swamps, where they accounted for 40.3% of all trees.
The number of well-drained forest type oligarchs recorded
in swamps was higher than expected by chance. For example,
if the 174 species that were recorded in both swamps and
the uplands were a random selection of the upland tree
ora, then they would be expected to include 26 of the
150 upland oligarchs. The observed number was 51
(chi-squared test, p 0.0001). The same pattern held for

Table 4. Forty-one tree species whose abundances and frequencies in a network of 25 1-ha tree plots at Yasun, Ecuador, suggest that they
are swamp specialists, swamp oligarchs, or both. Species followed by a hash sign (#) showed a signicant association with swamp forest in
an indicator species analysis. See the text for details.

Family
Arecaceae
Arecaceae
Arecaceae
Arecaceae
Arecaceae
Fabaceae
Arecaceae
Fabaceae
Elaeocarpaceae
Euphorbiaceae
Arecaceae
Polygonaceae
Arecaceae
Fabaceae
Rubiaceae
Malvaceae
Myristicaceae
Ochnaceae
Lauraceae
Combretaceae
Malvaceae
Urticaceae
Clusiaceae
Malvaceae
Lauraceae
Euphorbiaceae
Sapotaceae
Combretaceae
Rubiaceae
Annonaceae
Euphorbiaceae
Euphorbiaceae
Malvaceae
Moraceae
Euphorbiaceae
Fabaceae
Lecythidaceae
Sapindaceae
Clusiaceae
Fabaceae
Sapotaceae

Species

No. swamp
ind.

No. swamp
plots

Mauritiella armata
Mauritia exuosa#
Iriartea deltoidea
Euterpe precatoria#
Attalea butyracea
Macrolobium angustifolium#
Oenocarpus bataua
Zygia glomerata
Sloanea bark (NP3478)#
Hieronyma alchorneoides var. estipulosa#
Socratea exorrhiza
Triplaris weigeltiana
Astrocaryum murumuru var. urostachys
Pterocarpus amazonum/rohrii cf.
Isertia rosea#
Apeiba membranacea
Virola pavonis
Cespedesia spathulata
Licaria triandra#
Buchenavia amazonia#
Sterculia frondosa
Coussapoa trinervia#
Symphonia globulifera#
Theobroma subincanum
Lauraceae palermo (NP4484)
Sapium garcinia (NP2855)
Pouteria tenuipetiole (NP3323)
Buchenavia viridiora
Ferdinandusa guainiae
Unonopsis oribunda
Alchornea webster (NP4461)#
Euphorbiaceae nondescript (NP4465)
Sterculia colombiana
Ficus piresiana
Conceveiba 1 (NP1461)
Macrolobium acaciifolium
Couroupita guianensis
Sapindaceae chalky (NP4560)
Vismia myrsinac (NP1490)
Machaerium oribundum
Pouteria cuspidata ssp. robusta

378
359
343
327
146
131
76
54
52
45
44
39
30
30
28
21
21
21
20
18
18
17
16
12
11
10
9
8
8
7
7
7
7
7
6
6
6
6
5
5
5

1
4
3
5
2
5
3
3
4
4
4
3
3
4
4
4
3
2
4
4
3
4
4
3
1
2
3
1
1
3
3
1
3
4
3
2
3
2
2
3
3

oodplain forest oligarchs: 27 were expected in swamps and

50 observed (chi-squared test, p 0.0001).
Eight species qualied as oligarchs in all three forest types
(Table 4); the expected number under the null model was
less than one. These super-oligarchs accounted for 10.9,
13,9, and 26.5% of stems in upland, oodplain, and swamp
forests, respectively (Fig. 2, 4). On average, these species
occurred in 83.0% of plots in all forest types.
Species in swamps were more likely to be recorded in
oodplain forests than in upland forests, but that trend was
overshadowed by the much stronger trend towards species
occurring in all three forest types. Indeed, the number of species recorded in all three forest types was more than four times
higher than the number of species recorded in swamps and
oodplains but not in the uplands. Swamp species recorded
in the uplands but not in oodplains were very rare in swamps,

Oligarch?
Swamp
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1

Uplands

Floodplains

Specialist?
strict
strict

1
1

1
1
functional
functional

1
functional
strict

1
1

1
1
1
1

1
1

1
1

functional

strict

strict
functional
functional
1
strict
1

1
functional
functional

1
strict
strict

1
1
1
1
1

1
strict
strict
1

1
strict

1
functional
strict

1
1

while swamp species recorded in oodplains but not in

the uplands were relatively common in swamps (Fig. 3).

Discussion
While our data support all four of the well-documented
features of Amazonian swamps reviewed in the introduction
of this paper, they also indicate that those features coexist
side by side with strong and underappreciated patterns of
homogeneity. We begin this discussion by exploring four
relatively novel empirical patterns observed in swamps at
our study site. We then proceed to discuss how these four
patterns aect our understanding of the tree communities
growing in Yasun swamps. We close by highlighting key
questions and directions for further research.
909

Figure 4. The proportions of all species and stems surveyed in ve 1-ha swamp plots in Yasun, Ecuador, that belong to dierent
categories of oligarchic and non-oligarchic species. The category Oligarchs in all forest types consists of the eight species that are members
of the swamp oligarchy, the oodplain oligarchy, and the uplands oligarchy, while the category Oligarchs only in swamps consists of
the 14 species that are members of the swamp oligarchy but not members of the oodplain or uplands oligarchies.

Pattern 1: tree communities in swamps are

dominated by an oligarchy
The oligarchy of frequent and locally common tree species
observed in Yasun swamps (12.3% of swamp species
accounting for 59.0% of swamp stems) is similar in scale to
those observed in Yasun oodplains (17.2% spp., 55.2%
stems) and uplands (15.2% spp., 63.0% stems). Just as a
forester capable of identifying just 150 tree species in
the Yasun uplands could identify half of the standing
timber there (Pitman et al. 2001), the same forester would
need only 30 species to do the same in swamp forests.
Furthermore, this rule seems likely to apply across approximately 200 000 ha of swamp forest in Yasun National
Park. This represents additional evidence for the proposition that strong oligarchic components are a common
feature of Amazonian tree communities (Pitman et al.
2001; see also Burnham 2002, 2004, Pitman et al. 2003b,
2013, Vormisto et al. 2004, Maca and Svenning 2005,
Maca 2008, 2011, Fine et al. 2010).
While our data indicate that both the composition and
structure of tree communities growing on peaty substrates
in Yasun are predictable to some extent, they also make it
clear that there is nothing uniform about tree community
composition and structure in Yasun swamps. The fact that
only two of the 243 species recorded in swamps were
recorded in all ve swamp plots illustrates this very high
plot-to-plot variation. The ve plots we inventoried in
swamps are too small a sample to oer any understanding
of this swamp-to-swamp variation, which will in any case
require rigorous comparable data on variation in abiotic
conditions. In the meantime, our data suggest that the obvious swamp-to-swamp variation in composition and structure exists side by side with strong predictable components:
specically, a suite of species that are likely to be present in
910

a hectare of swamp forest regardless of whether the particular dominant in that forest is M. exuosa, I. deltoidea,
M. armata, or some other species.
For example, even the swamp plot with the most distinctive tree community (Dicaro 3, nearly half of whose stems
belong to a non-oligarch) contained 17 swamp oligarchs,
which together accounted for 38.6% of all stems. We thus
predict that the 30 swamp oligarchs identied here will
account for most stems in the majority of forests that
grow on peaty substrates in Yasun. This prediction is upheld
by data from Romero-Saltos et al.s (2001) inventory of
trees 10 cm dbh in ve 0.1-ha plots in Yasun swamps,
which found that 62.7% of stems belonged to nine species
identied here as swamp oligarchs.
By contrast, tree communities in many other swamps in
western Amazonia do not appear to be dominated by the
30 swamp oligarchs in Table 4. For example, the list of the
25 most abundant tree species in ve 0.1-ha swamp plots in
Caquet, Colombia, includes only three of our 30 swamp
oligarchs (Duque et al. 2001). The comparable number
for the 19 most abundant tree species in swamps at Pebas,
Peru, is four (Grndez et al. 2001). There is, however,
a small subset of our swamp oligarchs at Yasun that appear
to be abundant in swamps at all three sites (M. exuosa,
E. precatoria, and V. pavonis), and this suggests that there
may be a core group of tree species that are consistently
abundant in swamps at larger spatial scales.
Pattern 2: swamp specialists account for very few
species and a minority of stems in swamps
Of the 243 species recorded in swamps, 21 met our criteria
as swamp specialists. Together, these taxa represent slightly
less than a majority of stems in swamps (43.6%; Fig. 2), and

that proportion is lower when specialists are determined via

indicator species analysis (34.6%). Taken together with the
large degree of species-level overlap between swamps and
well-drained forest types (see next section), the tiny number
of swamp specialists ( 2% of all species recorded in our
Yasun tree inventory) suggests that evolutionary pressures
on trees to specialize on the specic environmental conditions in swamps have been rather weak over the several
million years that swamp forests have existed in the upper
Amazon (Hoorn et al. 2010).
The apparently tiny guild of swamp specialists we
observed also calls into question the commonly voiced
idea that habitat heterogeneity and its associated beta diversity inevitably increase gamma diversity on tropical
forest landscapes. If the 194 000 ha of swamp forests in
Yasun National Park really are contributing 30 unique
tree species to the regional tree ora, it is possible to argue
that the regional tree diversity would be higher if habitat
heterogeneity were lower, i.e. if there were no swamps on
the landscape. This is best illustrated by an extrapolation
from our data, which suggests that the two most common
swamp specialists (Mauritia exuosa and Mauritiella
armata) should have a combined population of 86 million
adult trees in the swamp forests of Yasun National Park.
It seems undeniable that if those stems were somehow
removed from swamps and added to Yasuns extremely
diverse and specialist-rich upland forests (i.e. by extending
the upland tree speciesarea curve with an additional 130
000 ha of uplands), they would contribute more than
two species to the regions gamma diversity. This is clearly a
simplistic extrapolation to a complex question, however,
and should not be mistaken for evidence that swamps
contribute little to regional diversity. It may be that swamps
in Yasun contribute few specialist trees but a large number
of specialist shrubs, vines, or herbs to the regional ora.
Alternatively, some of the 139 tree species we characterized
in this study as too rare or other may eventually be
shown to require swamps for their long-term persistence on
the landscape.
Finally, it is interesting to note that an unusually
large proportion of the Yasun swamp specialists listed in
Table 4 remain unidentied to species (4 taxa) or genus (3).
Most of these seven morphospecies have been matched
with fertile specimens that have been deposited in the
Missouri Botanical Garden for more than a decade, but a
recent review of their status conrmed that they are still
unidentied. This suggests that the small group of tree
species that specialize on swamps may feature a relatively
high rate of undescribed taxa. A similar pattern has been
observed for the broader swamp ora of Perus southern
Amazon (J. Janovec pers. comm.).

Pattern 3: Most tree species in swamps also occur

in adjacent well-drained forest types
Nearly all of the species we recorded in swamps (83.5%)
were also recorded in well-drained habitats. While this is not
a novel observation for Amazonian forests (see Pitman et al.
1999 for similar gures from southern Peru), it is much
more widespread than has been reported in the literature.

For example, while the pattern is apparent in the published

datasets of three other large-scale inventories in upper
Amazonia (Duivenvoorden et al. 2001), it was not reported
in the articles describing them (Duque et al. 2001, Grndez
et al. 2001, Romero-Saltos et al. 2001). Reviewing a dataset
collected in 30 0.1-ha plots in Caquet, Colombia, and
published as appendices by Duivenvoorden et al. (2001),
we found that 60.9% of tree species recorded in palm
swamps were also recorded in adjacent well-drained habitats.
The gures we calculated for data collected by the same
research group in Ampiyacu, Peru, and Yasun (a study independent of ours, but sharing some study sites) are 74.2 and
76.7% respectively (Duivenvoorden et al. 2001). For these
ve studies, spanning 11 degrees of latitude in Ecuador,
Peru, and Colombia, the mean proportion of swamp species
that were also recorded in adjacent well-drained forest types
was 76.0%. That most of the swamp tree ora at a site also
occurs in well-drained forest at the same site thus appears
to be a general feature of western Amazonian forests.
It is worth distinguishing this point from the observation
that well-drained habitats in Amazonia sometimes contain
small patches of poorly drained substrate where swamp
species may occur. While we have occasionally observed
Mauritia exuosa palms growing in small swampy patches on
upland terraces in western Amazonia, such patches were
not present in the well-drained plots we studied in Yasun
and are not the reason for the signicant compositional
overlap that we and others have documented between
swamps and well-drained forests.
The ve studies cited above also agree that the high
proportion of swamp species occurring in other forest
types is an underestimate, since the small number of plots
established at each site will under-sample the true number
of species that grow in multiple forest types there. In
both Caquet (Duque et al. 2001) and Yasun (RomeroSaltos et al. 2001), species-individual curves for woody
plants in palm swamps and well-drained forests did not
reach an obvious asymptote. Likewise, Romero-Saltos
et al.s (2001) inventory of 0.1-ha plots in Yasun swamps
recorded several species that are considered upland oligarchs in our study but which our own swamp plots
did not record. Thus, while our study in Yasun tallied 203
tree species that grow in both swamps and well-drained
habitats there, the true number is probably much higher.
And given that our inventory recorded a total of 596
species growing as adults in swamp and oodplain plots
at Yasun, it seems reasonable to suppose that the true
number of ood-tolerant tree species in the Amazon basin
is much higher than Junks (1989) estimate of 1000.
The signicant overlap in the tree oras of swampy
and well-drained habitats suggests that the ability to survive
on permanently waterlogged substrates is a feature shared
by a large segment of the upper Amazonian tree ora. This is
not surprising, considering the regions everwet climate;
even soils considered to be very well drained may in fact be
saturated for weeks on end during especially rainy periods.
This implies that all species in Yasun are adapted to
some extent to waterlogged soil conditions, which in turn
makes it unsurprising that some of the tree species common
in well-drained forest types in Yasun have been characterized as swamp specialists in drier regions of South America
911

(e.g. Tapirira guianensis, Richeria grandis, and Guarea

macrophylla in non-Amazonian Brazil, de Teixeira and
Assis 2011).

to which their seedlings in swamps are the product of

propagules from other forest types, the provenance of these
putative spillover populations in swamps must be considered unknown.

Pattern 4: many tree species in swamps are common

in well-drained forests

Synthesis and questions for future work

The tree species in swamps that are common in welldrained forests fall into two very dierent categories:
1) a small group of species that are common on both
waterlogged and well-drained soils, and 2) a large group of
species that are rare in swamps but common in other forest
types.
The rst group is typied by the eight super-oligarch
species, as well as the eight other swamp oligarchs that
are also oligarchs in either upland forests or oodplain forests (Table 4). This overlap of common species between
swamp, oodplain, and upland tree communities in Yasun
is most strikingly illustrated by the palm Iriartea deltoidea.
Although Iriartea is the most common tree in the regions
upland forests (where it averages 45 individuals ha1;
Pitman et al. 2001), its mean density is even higher in
swamps (69 ind. ha1), where it ranks as the third most
common species. Given that the species is also the most
common tree in oodplain forests (36 ind. ha1), the question of habitat preference in Iriartea deltoidea at large
spatial scales in Yasun seems moot. It would appear that the
species is not spilling over from a source to a sink, but is
simply outcompeting other species in dispersing to and
establishing in recruitment sites in all three forest types.
Svenning (1999) reported a similar result for I. deltoidea
in various microhabitats of a 50-ha upland plot at Yasun. In
a study of liana communities at Yasun, Burnham (2002)
likewise found one species to be the most common in both
upland and oodplain forests: Machaerium cuspidatum
(Fabaceae). It is a high priority to understand the mechanisms that allow these species to consistently outcompete
the other 1000 woody plant species in eastern Ecuador.
It would also be useful to know how long these superoligarchs have dominated Yasun s forests (i.e. whether an
adaptation that evolved relatively recently has allowed them
to become common by oering them a temporary escape
from natural enemies, or whether they have been outcompeting competitors for millennia).
The second and much more diverse group consists of
species that are relatively rare in swamps but common in
well-drained habitats. This group is typied by the 59 species that are oligarchs in well-drained habitats (and account
for 50% of stems there) but not oligarchs in swamps,
where they represent just 6.9% of stems. This very diverse
but numerically trivial component of the swamp community could reect occasional spillover into swamps of species
that prefer well-drained habitats (Shmida and Wilson 1985,
Rivas 2006). Given that these species survive to adulthood
in swamps, it should not be automatically assumed that
their tiny spillover populations there represent an ecological
sink (i.e. populations that would not persist without
the input of propagules from other forest types). Until additional studies can determine the extent to which these species are capable of regenerating in swamps, and the extent

The data presented here suggest that the conventional

picture of Amazonian swamp forests a forest type dominated by specialist tree species uniquely adapted to very wet
soil conditions is incomplete. We found that while specialists are unquestionably important in swamps, they are
accompanied there by two important non-specialist guilds:
one composed of species that are common and frequent
across both swamps and well-drained forests (superoligarchs) and the other composed of species that are
common in well-drained forests but occur only occasionally
in swamps (spillover). Indeed, these two non-specialist
guilds account for a much larger proportion of species and
a comparable proportion of stems in swamps as the two
specialist guilds (strict and functional specialists; Fig. 2).
These observations help bring our understanding of tree
communities in Amazonian swamps up to date with that of
wetland plant communities in the temperate zone, where
community composition and structure are commonly
understood to depend not merely on local environmental
conditions but also on the age, size, shape, and spatial
arrangement of swamp habitat (Horsk et al. 2012), on
the composition and diversity of adjacent, non-swamp
vegetation (Ashworth et al. 2006, Hettenbergerov and
Hjek 2011), and on the dispersal of propagules both
between swamps and from outside swamps (Houlahan et al.
2006). They also point to the somewhat counterintuitive
conclusion that the tree species composition and structure
of a given Amazonian swamp depend just as strongly on
the tree species composition and structure of the surrounding non-swamp forests as they do on the local guild
of swamp specialists. In turn, this helps explain Terborgh
and Andresens (1998) unexpected nding that ooded
forests in a given region of the Amazon are oristically
more similar to unooded forests in that same region than
they are to ooded forests in other regions.
Given our nding that a species likelihood of occurring
in a swamp increases with its abundance in non-swamp
forest types, seed dispersal in particular may play a much
more important role than previously appreciated in structuring Amazonian swamp communities. Very little is known
about dispersal processes within swamps and between
swamps and well-drained forest types, however, and how
they intersect with environmental lters in swamps. Much
of the data that is currently absent from the discussion
could be obtained by a combination of descriptive techniques (seed traps, phenological monitoring, seedling plots,
analyses of genetic structure in populations of species that
occur in both swamp and well-drained habitats) and experimental approaches (transplant and germination experiments) along transects extending from well-drained forest
into mixed and monodominant M. exuosa swamp forests.
If the transects in that study were arranged so as to allow
one to separate the eects of distance from well-drained

912

habitats from the eects of environmental conditions

expected to co-vary with that distance (e.g. peat depth,
ooding intensity), it could yield valuable insights regarding the degree to which environmental mechanisms and
dispersal are responsible for structuring tree communities in
tropical forests.
Acknowledgements We thank the following people for assistance
during the swamp portion of the eld work in Yasun: Humberto
Awa, Thomas Delinks, Edmundo Prez, Miles Silman, Wepera
Tocari, and Salvador. We are grateful for the hospitality shown to
our eld teams in the Huaorani communities of Dicaro and
Guiyero, and for institutional support during eld work from the
Yasun Scientic Station, the Tiputini Biodiversity Station,
the Ponticia Univ. Catlica del Ecuador, and the San Francisco
de Quito Univ. Rick Condit calculated Sorensons similarity
scores for all pairwise plot comparisons. Euridice Honorio and
Katy Roucoux kindly shared unpublished data from swamp
inventories in northern Peru.

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Supplementary material (Appendix ECOG-00774 at

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