Beruflich Dokumente
Kultur Dokumente
doi: 10.1111/ecog.00774
2014 The Authors. Ecography 2014 Nordic Society Oikos
Subject Editor: Jens-Christian Svenning. Accepted 21 February 2014
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.
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)
(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
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
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
Results
Structural and ecological attributes of the 25 tree plots are
summarized in Table 2, and additional details of the swamp
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.
170
108
44
44
11
11
10
10
10
7
Dicamumpare 2
ind.
68
36
28
27
23
19
18
17
12
12
378
95
71
64
18
13
11
11
10
10
Key
boldface top ten in multiple plots
palm
(S) swamp specialist
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.
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
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
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.
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
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
912
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