ECOGRAPHY 28: 515
/520, 2005

Abundance distribution relationships in fish assembly of the
/

Amazonas floodplain lakes

Carlos Granado-Lorencio, Carlos R. M. Araujo Lima$ and Javier Lobón-Cerviá

Granado-Lorencio, C., Araujo Lima C. R. M. and Lobón-Cerviá J. 2005.

Abundance
/ distribution relationships in fish assembly of the Amazonas floodplain
lakes.
/ Ecography 28: 515
/520.

This study shows that patterns in some community assemblages are not mainly
governed by local factors but also by regional ones. Using field data from 36 floodplain
forest lakes in the Amazon basin, we present transect count data on the richness,
abundance and distribution of floodplain lake fish species. A total of 194 fish species
were collected, of which 43 were classified as short-distance migrants. A positive
relation was found for local migratory and sedentary species abundances with
distribution at a regional scale. The study also suggests that the probability of the
presence of a migratory species is more affected by aspects of river-lake connectivity
than sedentary species. Our results seem to indicate that migratory species play an
important role in local dynamics of floodplain lakes.

C. Granado-Lorencio (granado@us.es), Dept of Biologı́a Vegetal y Ecologı́a, Univ. of

Sevilla, E-41080 Sevilla, Spain.
/ J. Lobón-Cerviá, Museo Nacional de Ciencias
Naturales, E-28006 Madrid, Spain.

Fish assemblage structure is not only determined by
local ecological mechanisms (competition, parasitism,
predation, and so on), but also by large scale environ-
mental factors (Ricklefs 1987, Menge and Olson 1990,
Loreau et al. 2001).
The Hot Spot Hypothesis (Diamond 1975) states that
a network of sites (within a region) are connected by
migrant species (dispersal). Therefore distribution of
each species will be the consequence of these ecological
adaptations of dispersal. In this scenario, migratory
species would have the role of connecting several
ecosystems temporarily, transferring matter and energy
(‘‘mobile link species’’, Myers 1993), at a regional scale
(sensu Hall 1972).
On the other hand, the most important contribution
of the metapopulation dynamics hypothesis was in
defining population structure in which local and regio-
nal-scale processes are linked (Levins 1969). This hy-
pothesis suggested that species with more extensive
distribution tend to be abundant locally; therefore, the
feature of a metapopulation is the need to adopt two
spatial scales to fully understand system dynamics: the
local patch scale and the regional patch network scale.
In the Amazon basin, there are four main freshwater
environments, the river, the lakes, the floodplains and the
tributaries. Approximately half of the total fish fauna are
in the tributaries. The dynamics of the river discharge are
the major force controlling Amazon landscape. The
water level varies from 8 to 12 m annually. Floodplains
lakes are areas that are very seasonal and are only
periodically inundated. Therefore, access by fishes de-
pends on the water level. Fish biomass of the Amazon
river is sustained, directly or indirectly, by material that
was produced inside of floodplain habitats (Junk et al.
1989).
Hanski et al. (1993) argue that the metapopulation
hypothesis cannot predict a relation between distribution
and abundance when migration is absent among local

$
deceased
Accepted 4 March 2005
Copyright # ECOGRAPHY 2005
ISSN 0906-7590

ECOGRAPHY 28:4 (2005) 515

communities. To test this argument, we investigated the
role of connectivity to inmigration for local community
structure. Hence foodplain lakes should be partly shaped
by regional factors due to this interconnectivity (open
community).
Methods
We chose a sector of 2000 km of the Amazon river,
between the Brazilian localities of Tefé and Parentins for
this study (Fig. 1). We sampled 36 forested floodplain
lakes that are connected to the river during the period of
maximum flood. The selection was based on: lake
surface area (20
/2000 ha) and connectivity (straight-
line distance between lake and river) (Table 1). The lakes
selected could be broadly classified in three main types
(Sippel et al. 1992): island lakes (12 lakes), 8 round lakes
(the lake has only a few major arms and minor shoreline
irregularities) and 16 dendritic lakes (the lake has a large
shoreline development). Diffuse connections with the
river channel was not considered.
We sampled fish using the same protocol during high
water season of 2000. Fish were collected using a set of
12 gill nets with 3
/11 cm mesh sizes (between opposite
knots). All nets were 25 m long and 2.5 m high. A new
set of nets was used per lake. Gill nets were set at the
edge of the flooded forest. We inspected the nets every
four hours during 24 h. To reduce seasonal variation of
NEGRO RIVER
AMAZONAS RIVER
21
19
17 20
6 15 18
9 11 13 16
5 7
12 14
3 4 8 10
2 Tefé
0 100 km
1 assemblages (see Table 1).
516
fish assemblages we used three independent boats and
teams, each one sampling one section of the basin (Tefé,
Coarı́ and Maués). All fish collected were preserved in
formaldehyde (4%) and transported to Inst. Nacional de
Pesquisas dâ Amazonia (INPA; Manaus, Brazil). Fishes
were classified as sedentary or short-distance migrant
species (sensu Böhning-Gaese and Oberrath 2003) based
on available regional information (Burgess 1989,
Barthem and Goulding 1997) (Table 2).
We defined a lake as the local scale, and defined a
region as the 36 lakes studied. We calculated local
species richness as the number of species found within
a single lake and regional richness as the number of
species found within the 36 lakes region. The distribution
was measured as the fraction of 36 lakes in which
the species was found, and local abundance as the size
of a local population based on relative abundances
(catch 24 h1).
We tested the effect of connectivity on migratory and
sedentary species with an analysis of co-variance with
linear regression and arc sine transform.
Results
A total of 196 fish species were collected (7695 fishes), of
which 43 species were classified as short-distance mi-
grants (Table 2). Except for highest and poorest lake
assemblages (species richness), the percentage of seden-
31
30
28 29 32
27 33
Santarém
24
Manaus
26 34
25 35
36
23
22
Fig. 1. Geographical
distributions of the floodplain
lakes used for analizing fish
ECOGRAPHY 28:4 (2005)
Table 1. Name, geographical co-ordinates and some characteristics of lakes studied.

Name Latitude Longitude Flood area Dry area Connection to river

(ha) (ha) length (km)

1 Tarará S 2825?21.0ƒ W 66830?10.0ƒ 167 121 0.4

2 Campina S 2836?40.0ƒ W 66820?55.0ƒ 233 17 1.8
3 Curimata S 2802?22.0ƒ W 66820?10.0ƒ 1889 311 2.1
4 Miriti S 1859?00.0ƒ W 66802?38.0ƒ 426 40 2.9
5 Sacambú S 2807?12.0ƒ W 66809?03.0ƒ 833 158 3.3
6 Tamaniquá S 2843?24.9ƒ W 65843?24.9ƒ 342 219 3.7
7 Malvado S 2819?33.0ƒ W 66821?03.0ƒ 65 55 8.8
8 Uará S 2842?39.6ƒ W 65836?15.6ƒ 154 120 4.7
9 Tracajá S 3806?08.5ƒ W 64846?08.8ƒ 151 35 1.3
10 Alvaraes S 3812?07.8ƒ W 64850?00.8ƒ 145 77 2.7
11 sin nombre S 3853?29.4ƒ W 62835?11.4ƒ 108 28 0.0
12 Ipixuma S 3851?16.2ƒ W 63852?39.6ƒ 479 345 1.8
13 Cipotuba S 3849?19.2ƒ W 62820?54.0ƒ 272 157 1.8
14 Anori S 3845?03.4ƒ W 61841?03.6ƒ 594 236 7.6
15 Urucurı́ Grande S 3855?28.8ƒ W 62800?16.8ƒ 683 674 3.2
16 Sao Tomé Grande S 3855?21.0ƒ W 61828?36.6ƒ 733 194 2.7
17 Rasgado S 3836?25.2ƒ W 61809?30.0ƒ 142 83 1.3
18 Jacaré S 3838?35.4ƒ W 60850?06.0ƒ 783 103 6.1
19 Acuru S 3836?30.0ƒ W 63841?48.0ƒ 315 0 4.3
20 Comprido S 3845?05.4ƒ W 63832?31.8ƒ 559 96 4.9
21 Apaurá S 3854?18.6ƒ W 62825 57.6ƒ 1031 737 0.7
22 Ajura S 3853?29.4ƒ W 62835?11.4ƒ 171 153 2.0
23 Luis S 3854?21.0ƒ W 62848?09.0ƒ 137 93 1.1
24 Estasio S 2848?02.8ƒ W 57813?24.5ƒ 607 392 1.6
25 Carar-açú S 2821?55.7ƒ W 57835?43.4ƒ 388 77 5.0
26 Terra Preta S 2826?52.0ƒ W 57838?13.4ƒ 1444 296 1.5
27 Apunuma S 2854?02.0ƒ W 58805?53.6ƒ 327 138 0.4
28 Piloto S 2831?80.9ƒ W 57812?43.4ƒ 920 176 0.8
29 Arari S 2830?81.9ƒ W 57811?44.2ƒ 1088 825 0.5
30 Coró-coró S 3813?55.5ƒ W 58841?28.7ƒ 14 10 4.6
31 Moaná S 2852?37.0ƒ W 57831?53.0ƒ 1392 1156 1.4
32 Taperebá S 3818?35.9ƒ W 58840?01.0ƒ 755 199 2.6
33 Garças S 3810?30.9ƒ W 57842?13.0ƒ 408 371 4.2
34 Paracuuba S 3807?30.6ƒ W 59816?26.1ƒ 47 13 8.6
35 Arroz S 2846?35.9ƒ W 57859?39.5ƒ 673 33 2.8
36 Buiuçú S 3803?42.9ƒ W 57844?47.0ƒ 1333 828 2.3

tary species showed no significant pattern. A positive Discussion

relation between local sedentary and migratory richness
was observed where local species richness increased with
the number of migratory and sedentary species, and
sedentary species were always dominant (F /28.47; pB/
0.01; DF /1,32, Fig. 2). The proportion of ‘‘migrants’’
was similar for all local populations (p /0.654, Fisher
test). A significant positive relation did exist for propor-
tion of migrants for both average abundance and
distribution. The regression is significant (sedentary:
F/38.230; DF /151; pB/0.001; R2 /0.21; migratory:
F/105.03; DF/43; pB/0.001; R2 /0.26; Fig. 3). No
significant relation of the residual on distribution but,
migratory species had, generally, positive residuals in
abundance-distribution whereas sedentary species
showed no tendencies (Fig. 4).
The lake-river connectivity on migratory or sedentary
species richness chart the role of the seasonal focus
(river) on the lake assemblages. We found that only has
been awkward significant for migratory species (Fig. 5).
Migrating species are more affected by connectivity than
sedentary species (F /11.93; DF /1; pB/0.001; R2 /
0.27).
Most of the empirical research in metapopulation
ecology has focused on terrestrial and marine systems,
while studies in freshwater systems are scarce. From a
metapopulation perspective, our results corroborate
some previous findings of abundance-distribution rela-
tion. Numerous studies have suggested that a relation
exists between distance to settlement source and local
richness (Cornell and Lawton 1992, Osborne and Wiley
1992, Detembeck et al. 1992, Hugueny and Paugy 1995).
The average abundance
/ distribution relation may
result from metapopulation dynamics but also from
sampling bias (Wright 1991); rare species, or species with
low capturability have a higher probability to be absent
from a sample than abundant species.
Distance between lake and river is a major factor of
connectivity where migrating species are more affected
by connectivity than sedentary species. Connectivity was
expected to be negatively affected by distance between
river and lake, as generally assumed (Porter and Dooley
1993, Moilanen et al. 1998). We found that the lake-river
connection is a homogeneity factor of local fish assem-
blages, strongly associated to migratory regional richness

ECOGRAPHY 28:4 (2005) 517

Table 2. Fish species collected during all sampling lakes. Names in bold-faced represent migratory fish species.

Acarichthys heckelii Curimatella meyeri Nemadoras hemipeltis Sternarchella orthos

Acaronia nassa
Acestrorhynchus falcatus
Acestrorhynchus falcirostris
Acestrorhynchus heterolepis
Acestrorhynchus microlepis
Aequidens sp.
Agamyxis pectinifrons
Ageneiosus atronasus
Ageneiosus brevifilis
Ageneiosus brevis
Ageneiosus sp.
Ageneiosus ucayalensis
Agoniates anchovia (M)
Amblydoras affinis
Anadoras weddellii
Ancistrus dolichopterus
Anodus elongatus (M)
Anodus orinocensis (M)
Apteronotus albifrons
Apteronotus hasemani
Arapaima gigas
Argonectes longiceps (M)
Astronotus crassipinnis
Astronotus ocellatus
Astyanax sp.
Auchenipterichthys thoracatus
Auchenipterus ambyiacus
Auchenipterus britskii
Auchenipterus nuchalis
Boulengerella lucia
Boulengerella maculata
Brycon cephalus (M)
Brycon melanopterus (M)
Bryconops alburnoides
Bryconops giacopinii
Callichthys callichthys
Calophysus macropterus
Catoprion mento
Centromochlus heckelii
Chaetobranchus flavescens
Chaetobranchus semifascia-
tum
Chalceus erythrurus
Charax sp.
Cichla monoculus
Cichla temensis
Cichlasoma amazonarum
Colomesus asellus
Colossoma macropomum (M)
Crenicichla joana
Crenicichla lugubris
Crenicichla reticulata
Crenicichla sp.
Ctenobrycon hauxwellianus
Curimata inornata (M)
Curimata knerii (M)
Curimata ocellata
Curimata vittata
Curimatella alburna
Cynodon gibbus
Cyphocharax abramoides
Dekeyseria amazonica
Dekeyseria scaphirhyncha
Dianema longibarbis
Dianema urostriatum
Eigenmannia limbata
Eigenmannia sp.
Eigenmannia virescens
Geophagus altifrons
Geophagus proximus
Glyptoperichthys gibbiceps
Gymnotus carapo
Hemiodus argenteus (M)
Hemiodus atranalis
Hemiodus immaculatus (M)
Hemiodus microlepis (M)
Hemiodus sp.1
Hemiodus ternetzi
Hemiodus unimaculatus (M)
Heros sp.
Hoplerythrinus unitaeniatus
Hoplias malabaricus
Hoplosternum littoralis
Hydrolycus scomberoides
Hypophthalmus edentatus (M)
Hypophthalmus fimbriatus (M)
Hypophthalmus marginatus (M)
Hypoptopoma gulare
Hypostomus hoplonites
Hypostomus sp.1
Hypostomus sp.2
Hypselecara temporalis
Ilisha amazonica
Laemolyta proximus
Laemolyta taeniatus
Leiarius pictus
Leporinus affinis
Leporinus amazonanum
Leporinus fasciatus (M)
Leporinus friderici
Leporinus trifasciatus (M)
Liposarcus pardalis
Loricariichthys acutus
Loricariichthys maculatus
Loricariichthys nudirostris
Lycengraulis grossidens
Megaladoras uranoscopus
Megalechis thoracatum
Mesonauta insignis
Metynnis argenteus
Metynnis hypsauchen
Metynnis luna
Metynnis maculatus
Moenkhausia grandisquamis
Myleus rubripinnis
Mylossoma aureum (M)
Mylossoma duriventre (M)
Nemadoras humeralis Sternarchella schotti
Opsodoras cf. stuebelii Sternopygus macrurus
Osteoglossum bicirrhosum Sternopygus obtusirostris
Oxydoras niger Stethaprion erythrops
Pachypops trifilis Stichonodon insignis
Parauchenipterus galeatus Symphysodon aequifasciata
Paulicea luetkeni (M) Tatia intermedia
Pellona castelnaeana (M) Tatia sp.
Pellona flavipinnis (M) Tetragonopterus chalceus
Piaractus brachypomus (M) Thoracocharax securis
Pimellodina flavipinnis (M) Thoracocharax stellatus
Pimelodus blochii Trachelyopterichthys taeniatus
Pirinampus pirinampu (M) Triportheus albus (M)
Plagioscion montei Triportheus culter
Plagioscion squamosissimus Triportheus elongatus (M)
Potamorhina altamazonica (M) Triportheus flavus (M)
Potamorhina latior (M) Uaru amphiacanthoides
Potamorhina pristigaster
Pristigaster cayana
Pristobrycon serrulatus
Pristobrycon sp.
Prochilodus nigricans (M)
Psectrogaster amazonica (M)
Psectrogaster rutiloides (M)
Pseudanos gracilis
Pseudanos trimaculatus
Pseudoplatystoma fasciatum (M)
Pseudoplatystoma tigrinum (M)
Pseudorinelepis genibarbis
Pterodoras lentiginosus
Pterophyllum scalare
Pygocentrus nattereri (M)
Pygopristis denticulatus
Raphiodon vulpinus (M)
Rhabdolichops caviceps
Rhabidolichphs caviceps
Rhamphichthys marmoratus
Rhamphichthys sp.
Rhytiodus argenteofuscus (M)
Rhytiodus microlepis (M)
Roeboides myersi
Satanoperca acuticeps
Satanoperca daemon
Satanoperca jurupari
Satanoperca lilith
Schizodon fasciatum (M)
Semaprochilodus insignis (M)
Semaprochilodus taeniurus (M)
Serrasalmus altispinis
Serrasalmus compressus
Serrasalmus eigenmanni
Serrasalmus elongatus
Serrasalmus manueli
Serrasalmus rhombeus
Serrasalmus sp.1
Serrasalmus sp.2 (robertsoni )
Serrasalmus spilopleura
Sorubim lima

(Huston 1997, Meschiatti et al. 2000). Undoubtedly,
many physico-chemical and ecological factors are likely
to vary in floodplain lakes as distance from river
increases and the fact that dispersal is not obligatorily
are the most plausible explanation for the observed
pattern.
Our data support the arguments of Fahrig and
Merriam (1994) that the probability of recolonisation
depends on (among others): dispersal characteristics of
the migrant species and temporal changes in landscape
structure. In the lower water period, a lake is more
isolated and its volume is reduced. If the lake doesn’t dry
out completely, the co-existing species compete strongly
until the following rainy period. At the regional scale,
the local assemblages (set of species) would be consid-
ered as open communities or belonging to metacomu-
nities (communities organized into networks linked by
dispersal; Wilson 1992, McGill 2003).

518 ECOGRAPHY 28:4 (2005)

 sedentary 3
70
Sedentary and migratory

migratory

60 y = 0.6428x + 1.8302
2 2

Average abundance (residual)

R = 0.9412
50
richness

40 1
30
20 0
y = 0.3096x - 0.0726
2
10 R = 0.7674
0 -1
0 50 100
Local richness
-2
Fig. 2. The relationship between local richness (number of 0.0 0.2 0.4 0.6 0.8 1.0
species/lake) and migratory species richness (number of migra- Distribution
tory species/lake) for all lakes studied.
Fig. 4. Average abundance (residual)
/ distribution relation of
migratory (closed circle) and sedentary species (open circle).
Little is known about colonization processes in Each point represents one species.
tropical aquatic ecosystems. According to our results
(e.g. migratory species richness), abundance in a geo- lakes. However, we stress that this work is preliminary
graphical distribution and lake-river connectivity (isola- and much more data are needed before conclusions can
tion degree) plays a role in local dynamics of floodplain be drawn about the fish assemblages of floodplain lakes
as a metapopulation (interconnected populations that
function as a unit). Several questions have become major
y = 0.768x + 0.3054
1.6 2 investigations. Our contribution, in an integrated view of
R = 0.2694
1.4
Amazon fishes, is one more example that demonstrates
Log10 average abundance

that fish community assembly (Drake 1990, Morton and

1.2 Law 1997) is not only influenced by local mechanisms,
1 but also by processes acting on much larger spatial scales
(Hubbell 2001, Wootton 2001).
0.8

0.6 60
Sedentary local richness

y = -1.3024x + 32.32
2
0.4 50 R = 0.1185

0.2 40

0 30
0 0.2 0.4 0.6 0.8 1 20
Distribution
10
1.6 y = 0.8021x + 0.2116
2
0
R = 0.2106 0 2 4 6 8 10 12
Log 10 average abundance

1.4
Connectivity
1.2
25 y = -0.3328x + 18.768
Migratory local richness

1 2
R = 0.5095
0.8 20

0.6 15

0.4 10
0.2
5
0
0
0 0.2 0.4 0.6 0.8 1 0 10 20 30 40
Distribution Connectivity

Fig. 3. The relationship between average abundance (fish Fig. 5. Relationship between connectivity (straight-line distance
24 h 1) and distribution for migratory (top) and sedentary between lake and river) and local migratory and sedentary
species (bottom). species.

ECOGRAPHY 28:4 (2005) 519

Acknowledgements
/ This study was supported by Agencia
Española de Cooperación Internacional (AECI, España) and
Inst. Nacional de Pesquisas dâ Amazonia, Manaus, Brasil. The
assistance of the technical staff at the Inst. Nacional de
Pesquisas dâ Amazonia, Manaus, Brasil, is gratefully
acknowledged. A. Klemetsen (Subject Editor) provided several
insightful comments and criticisms on earlier versions of the
manuscript.
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