Received: 30 August 2023 Accepted: 28 November 2023

DOI: 10.1111/icad.12707

ORIGINAL ARTICLE

It is not a mere piece of the mainland: Fluvial islands contribute

to regional spider diversity in a seasonally flooded Amazonian
archipelago

Thiago Gomes de Carvalho 1 | Thierry Ray Gasnier 2 | Sérgio Henrique Borges 1,2

1
Programa de Po
s-Graduação em Zoologia,

gicas, Universidade
Instituto de Ciências Biolo
Federal do Amazonas (UFAM), Manaus, Brazil
2
Departamento de Biologia, Universidade
Federal do Amazonas (UFAM), Manaus, Brazil
Correspondence
Sérgio Henrique Borges, Programa de Po
s-
Graduação em Zoologia, Instituto de Ciências

gicas, Universidade Federal do Amazonas
Biolo
(UFAM), Manaus, Brazil.
Email: shborges9@gmail.com
Funding information
Fundação de Amparo à Pesquisa do Estado do
Amazonas; Programa Fixam, Grant/Award
Number: 017/2014; Universal Amazonas,
Grant/Award Number: 002-2018
Editor: Yves Basset and Associate Editor:
James Bell
Abstract
1. Islands and analogous insular habitats are found in several ecological contexts. With
the exception of oceanic islands, however, the contribution of these insular habitats
to regional biodiversity remains poorly investigated, especially in tropical regions.
We investigated the understory spider assemblages in an Amazonian river archipel-
ago to evaluate the contribution of fluvial islands to regional spider diversity.
2. We collected spiders on 30 islands ranging from 1 to 1527 ha and at 23 sites in

flooded forests and terra firme
mainland habitats represented by continuous igapo
forests. Spiders were collected by shaking the vegetation over a white sheet during
a standardised time of 30 min.
3. From 1404 adult individuals, we identified 290 spider species or morphospecies,

flooded for-
with 150 species being recorded on the fluvial islands, 122 in the igapo
ests and 103 in the terra firme forests. Spider assemblages of fluvial islands exhibit
lower alpha diversity and equitability compared with mainland habitats. The species
composition of fluvial islands was significantly distinct from that of mainland habi-

flooded forests. Several families
tats, even when compared with neighbouring igapo
and species of spider were significantly associated with a particular habitat, includ-
ing 8 families and 10 species identified as indicators of fluvial islands.
4. Fluvial islands effectively contribute to regional diversity since their spider fauna is
not a mere impoverished subset of species from mainland habitats. Events that alter
the flood pulse in Amazonian rivers (e.g., construction of hydroelectric dams) could
threaten the unique biological assemblages of the floodplain habitats, especially the
fluvial islands.

KEYWORDS
alpha diversity, Amazonian arthropods, dominance, flooded forests

I N T R O D U CT I O N Whittaker & Fernández-Palacios, 2009). In addition, islands have high

Biological assemblages on islands diverge markedly from those found
in mainland continuous habitats with equivalent size by presenting
lower species diversity, higher population densities and distinct spe-
cies composition (Adler & Levins, 1994; MacArthur & Wilson, 1967;
levels of endemism and are more susceptible to biological invasions
and species extinctions (Kier et al., 2009; Moser et al., 2018).
Although most documented patterns of biological diversity on islands
were established from oceanic islands and archipelagos (Whittaker &
Fernández-Palacios, 2009), islands and analogous insular habitats exist

Insect Conserv Divers. 2023;1–11. wileyonlinelibrary.com/journal/icad © 2023 Royal Entomological Society. 1


2 DE
in several ecological contexts around the planet, especially in tropical
regions. However, our understanding of the contribution of these
islands to regional biodiversity remains insufficient (Itescu, 2019;
Prance, 1996).
The Amazon rainforest is the largest tropical forest on the planet
and its remarkable biological diversity is intrinsically associated with
past geological events and vegetation heterogeneity (Dambros
et al., 2020; Figueiredo et al., 2017; Haffer, 2001). Even though most
of the lowland in the Amazon is essentially covered by continuous
upland forests, there are numerous true islands and island-like habi-
tats in the region, such as mountaintops, savanna enclaves and
patches of vegetation growing in white-sand soils (Costa et al., 2020;
Giulietti et al., 2019; Magnusson et al., 2008). Fluvial islands found in
the channels of major Amazonian rivers are among these insular
habitats.
Islands are conspicuous features of the Amazon riverine land-
scapes and, like other habitats in the floodplains, they are subject to
seasonal flooding (Junk et al., 2011). Fluvial islands experience a dra-
matic reduction of their areas due to seasonal flooding that can range
from 6 to 16 m in height, with some sites remaining underwater from
4 to 6 months every year (Junk et al., 2011). Thus, seasonal flooding
acts as a strong environmental filter by selecting a peculiar set of ani-
mal and plant species found in the floodplain habitats, including on
fluvial islands (Arias et al., 2018).
Previous studies suggest that some animal species are specialised
in the use of fluvial islands, therefore making their assemblages dis-
tinct from the other habitats (Borges et al., 2019; Laranjeiras
et al., 2021; Nunes et al., 2014). Rosenberg (1990), for example, found
that at least 18 bird species are specialised in the use of fluvial islands
in the western Amazon. Documented evidences of distinct fauna
composition on fluvial islands can be relevant for regional conserva-
tion strategies, because islands are disproportionally susceptible to
environmental impacts when compared with continuous habitats due
to their spatial restriction and small occupied area. The contribution of
Amazonian river islands to regional diversity, however, has been
assessed on only a few occasions and covered a limited range of taxo-
nomic groups (Laranjeiras et al., 2021; Nunes et al., 2014;
Rosenberg, 1990).
The use of Amazonian fluvial islands by spiders has not been
investigated. Spiders are among the most species-rich predatory
arthropods and are widely distributed across the planet’s ecosystems
(Bonaldo et al. 2009a; Brescovit et al., 2002; Coddington &
Levi, 1991). Spiders are particularly useful for investigating diversity
patterns on islands due their ability to disperse on land, water and
even through the air (Carvalho & Cardoso, 2014; Gillespie, 2002;
Kuntner & Agnarsson, 2011; Suter et al., 2003; Vanbergen
et al., 2017). Unfortunately, the scarcity of ecological studies makes it
difficult to establish the contribution of major habitats to Amazonian
spider diversity (Bonaldo et al. 2009a; Coddington et al., 2009;
Höfer, 1990; Rego et al., 2009).
Here, we present a quantitative analysis of the spider assem-
blages of fluvial islands in the Anavilhanas archipelago, one of the larg-
est networks of river islands in the world (Latrubesse &
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CARVALHO ET AL.
Stevaux, 2015). The main goal of the study is to evaluate the contribu-
tion of fluvial islands to regional spider diversity. To achieve this aim,
we sampled spiders on fluvial islands and in mainland habitats repre-

flooded
sented by flooded forests that occupy the river margins (igapo
forests) and upland terra firme forests. In addition, we performed a
brief analysis of the major anthropogenic impacts on river channels
that could threaten the biological assemblages of Amazonian fluvial
islands.
We used the general biotic patterns identified in island biogeogra-
phy studies (Whittaker & Fernández-Palacios, 2009) as a framework
to establish the following predictions: (i) considering the area-effects
on island biological diversity (MacArthur & Wilson, 1967; Triantis
et al., 2012), it is expected that mainland habitats, which occupies
larger and continuous areas, will have a greater spider alpha diversity
than fluvial islands yet that these habitats are sampled with similar
efforts; (ii) fluvial islands present a certain degree of isolation from
mainland habitats and experience dramatic seasonal flooding, which
reduce habitat area and resources availability. Under these ecological
conditions, species possessing greater dispersal abilities and better
adapted to flooding (Adis & Junk, 2002; Höfer, 1990) are more likely
to successfully colonise the islands and become more abundant. Thus,
it is expected that spider assemblages in fluvial islands will be domi-
nated by few abundant species (low equitability) compared with main-
land habitats; (iii) the islands in the studied system are relatively close
to mainland habitats and, therefore, it is expected a high overlap in
species composition between islands and non-island habitats, espe-

flooded forests that occupy river mar-
cially from neighbouring igapo
gins and had vegetation structure and floristics similar to islands;
(iv) based in the previous results obtained with other taxa such as bee-
tles (Nunes et al., 2014) and birds (Rosenberg, 1990), we hypothesise
that some spider species might be specially adapted to live on fluvial
islands and their frequency and abundance will be likely higher in this
habitat.
M A T E R I A L S A N D M ET H O D S
Study area
Spider sampling took place in the lower course of the Negro River
within the limits of the Anavilhanas National Park (350,000 ha), which
is a federal conservation area created to protect one of the largest flu-
vial archipelagos in the world (ICMBio, 2017). The vegetation of the

flooded forest with tree heights vary-
studied islands consists of igapo
ing from 15 to 20 m (Piedade et al., 2005). The islands selected for
the study have narrow shapes and variable sizes (Figure 1).
The studied fluvial islands are embedded in a dynamic and hetero-
geneous landscape that is seasonally inundated (Junk et al., 2011;
Latrubesse & Stevaux, 2015). Differences between the maximum and
minimum levels of the Negro River are on average 11 m, and this sea-
sonal dynamic in the river level drastically alters the regional land-
scapes, though even more so on the islands (ICMBio, 2017; Junk
et al., 2011). During maximum high-water period (from May to July),

SPIDERS IN Amazonian FLUVIAL ISLANDS
F I G U R E 1 Study region highlighting in green the islands where the spiders were collected. Stars represent collecting sites on islands and

flooded forests and terra firme forests). The darker grey line indicates the partial limits of the Anavilhanas
triangles in mainland habitats (igapo
National Park.
much of the islands’ vegetation is submerged, leaving only the cano-
pies of the tallest trees exposed (Almeida-Filho et al., 2016;
Latrubesse & Stevaux, 2015). In the low-water period (from
September to December), the islands fully emerged and sand and clay
banks are exposed on the riverbed, thus promoting terrestrial connec-
tivity between some islands (Almeida-Filho et al., 2016;
ICMBio, 2017).
In addition to fluvial islands, we sampled spiders in continuous

flooded forests and the terra firme
habitats represented by the igapo
forests that are localised on the margins of the Negro River (Figure 1).

forests are seasonally flooded like the islands
The continuous igapo
and occupy the banks of the Negro River and its tributaries. In con-
trast, terra firme forests occupy the highest parts of the terrain and
never get flooded. Even though they are connected, igap

o and terra
firme forests show marked differences in vegetation structure and flo-
ristic composition (Quaresma et al., 2018).
Spider sampling and identification
Spiders were sampled in September 2017 and December 2018, which
are the period of the low-water season when the islands fully
emerged. The spiders were collected using an adaptation of the vege-
tation beat method (Harris et al., 1972), with a white sheet measuring
1  1.5 m positioned under the vegetation in the understory of the
studied habitats. The shrubs, bushes and small tree branches above
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3
the sheet were vigorously shaken, and the spiders that fell on the
cloth were collected and placed in containers with 90% alcohol. Col-
lecting effort in each sampling site was around 30 min, and the same
field protocol was applied in the island and mainland habitats. We
searched for sampling spots as similar as possible in vegetation struc-
ture and with a similar number of shrubs and small trees in the islands
and mainland sites. Spider collection in the Anavilhanas National Park
was authorised by the Brazilian Minister of Environment through the
Sistema de Autorização e Informação em Biodiversidade (Sisbio) under
the licence 58392-2.
Anavilhanas harbour 100 islands with distinct sizes, shapes and
variability in vegetation structure. To collect spiders in such heteroge-
neous landscapes, the islands were selected using the following cri-
teria: (i) variability in size, (ii) proximity to the left and right of the
Negro River margins and (iii) facility to access and collected spider in
several islands in a field day. The sizes of islands were estimated using
Google Earth Pro by drawing a polygon around the island on a satellite
image from December 2016. We have sampled spiders in 30 islands
with highly variable area (mean = 352.6 ha, 1–1523 ha, coefficient of
variation = 136.4%) and well distributed in the core area of the Ana-
vilhanas archipelago (Figure 1). Here, we are interested in quantifying
the biological distinctness of the islands compared with other habitats,
thus detailed information on islands attributes and analysis on the
effects of island size on spider diversity will be presented and
explored in a separate study. In addition, we made spider collections
at 13 sampling sites in igap

o flooded forests and at 10 sites in terra

4 DE
firme forests. Each collection point was treated as a sampling unit for
the analysis, thus totalizing 53 sample units, 30 on fluvial islands and
23 in mainland habitats.
The spiders were identified to the most refined taxonomic rank
possible using keys for spider families (Brescovit et al., 2002; Ubick
et al., 2005) and reviews for the genera, available on the websites: The
World Spider catalogue (http://www.wsc.nmbe.ch), The Peckham
Society (http://peckhamia.com/salticidae), Jumping spiders (Arachnida:
Araneae: Salticidae) of the world (http://www.jumping-spiders.com) and
Araneae (https://araneae.unibe.ch/key). Males and females were
included in species count. Most species had at least a few individuals
collected from the same location, which helped paring them. In a small
number of cases where individuals did not match any species, they were
treated as a single species. All identified individuals were incorporated
into the Paulo Bührnheim Zoological Collection at the Federal Univer-
sity of Amazonas (CZPB–UFAM).
Most individuals were identified to family level, but only the
adults could be safely assigned to species or morphospecies. High-
quality photographs of living adults were used to characterise the spe-
cies or morphospecies. The spiders chosen to represent the species
(photographic type) were individuals with all limbs and without defor-
mities. The photographs produced during this study will be used in a
further publication of an illustrated inventory of spiders of the Anavil-
hanas archipelago using the same approach as Carvalho and Gas-
nier (2019).
Analyses
To compare spider alpha diversity from the three habitats, we built
individual cumulative curves extrapolated for a common sampling
effort and coverage-based curves (Chao & Jost, 2012; Colwell
et al., 2012) using the INEXT freeware (https://chao.shinyapps.io/
iNEXTOnline/). We extrapolate the samples to 1000 individuals in
each habitat using the method of Colwell et al. (2012). Sampling cov-
erage (SC) in each habitat was estimated using the formula presented
in the study by Chao and Jost (2012) that uses three parameters:
number of singletons (species represented by 1 individual or f1), num-
ber of doubletons (species represented by 2 individuals or f2) and the
total number of individuals in the sample (n). The non-overlap of
the confidence intervals in the curves indicates significant differences
in the estimated species diversity between habitats (Chao &
Jost, 2012, Colwell et al., 2012).
We quantified alpha diversity using the Hill series, which inte-
grates distinct aspects of relative abundance and species richness
(Hill, 1973). In this approach are calculate indices using different coef-
ficients (q) that emphasise rare species (q = 0, species richness), nei-
ther rare nor common species (q = 1, exponential of Shannon’s
entropy), and only the most abundant species (q = 2, inverse of Simp-
son’s concentration index) (Chao et al., 2014; Roswell et al., 2021).
The formulas of mentioned indices and description of the sampling
standardisation can be found in studies by Colwell et al. (2012), Chao
and Jost (2012), Chao et al. (2014) and Roswell et al. (2021). In
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CARVALHO ET AL.
complement, we have measured equitability of the samples using Pie-
lou’s evenness indices (Magurran, 2004) and tested the differences
between habitats through a Krusal–Wallis test with Dunn contrast.
Non-metric multidimensional scale (NMDS) analyses were used
to represent variations in species composition (i.e., beta diversity).
These ordination analyses were performed with abundance data
(Bray–Curtis distance) relativized using the total number of individuals
collected in each sample and square root transformed to equalise the
contribution of abundant and rare species (Clarke & Warwick, 2001).
NMDS ordinations were also run with presence or absence data
(Jaccard distance). Compositional analyses were performed with all
species or morphospecies as well as only the most abundant ones
(≥10 individuals collected). Analyses of similarity (ANOSIM) were run
to test the hypothesis of there being differences in species composi-
tion between habitats. Ordinations and ANOSIM analyses were per-
formed using the vegan packages and Primer v6 (Clarke &
Gorley, 2006; Oksanen et al., 2019).
An indicator species analysis (De Cáceres & Legendre, 2009) was
performed to verify whether some taxa are significantly associated
with a particular habitat. In this analysis, relative abundance and fre-
quency of species are used to calculate an indicator value (IndVal)
ranging from 0 to 1 according to the degree of association of a species
to a particular habitat with statistical inferences based on 9999 per-
mutation tests (De Cáceres & Legendre, 2009).
As commonly occurs in spider inventories, most collected individ-
uals were juveniles, thus precluding an accurate identification at spe-
cies level. In contrast, most individuals could be reliably assigned to
family level. Then, for a better use of the data, we run indicator spe-
cies analysis at family and species resolutions. Only families with more
than 50 individuals and species with more than 10 individuals were
included in these analyses.
RE SU LT S
General inventories results
A total of 4905 spiders were collected, which comprised 3501 juve-
niles and 1404 adults (Supporting Information S1). Of this total, 3278

flooded forests
spiders were collected on fluvial islands, 942 in igapo
and 685 in terra firme forests. We recorded 38 spider families, with
29 on fluvial islands, 29 in igap

o flooded forests and 32 in terra firme
forests.
We identified 290 species or morphospecies among the 1404

flooded
adult individuals, 150 species on fluvial islands, 122 in igapo
forests and 103 in terra firme forests. About 20% of the adults
belong to the species Scopocira aff. carinata, the most abundant and
frequent spider in the samples. Only 11 species were present in all
inventoried habitats. A substantial portion of species and morphos-
pecies (46%) was represented by a single individual (singletons) in
the samples. The proportion of these singletons was higher in terra
firme forests (44%), followed by igap

o flooded forests (35%) and flu-
vial islands (31%).
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SPIDERS IN Amazonian FLUVIAL ISLANDS 5

F I G U R E 2 Patterns of alpha diversity of spiders in Anavilhanas National Park using equal sampling-effort (a–c) and sampling coverage (d–f)
standardizations applied to Hill numbers (see Analysis section); 95% confidence intervals resulting from 100 permutations are shown in different
coloured shapes. Vertical dashed lines in graphs d–f indicate sampling coverage similar between the studied habitats.

F I G U R E 3 Ordinations of sampling sites by non-metric multidimensional analysis (NMDS) using the Bray–Curtis distance with relative
abundance data. Analysis includes all spider species/morphospecies (a) or only those considered abundant (≥10 individuals—b).

Spider assemblages: Alpha and beta diversities
Species richness per sample was similar between habitats: igap

o flooded
forests (average: 14 species, 7–20 species), terra firme forests (average:
17 species, 10–28 species) and fluvial islands (average: 15 species, 8–22
species). As expected, SC was more effective in islands (SC = 0.92,

flooded forests (SC = 0.75, f1 = 70,
f1 = 66, f2 = 27) than igapo
f2 = 18) and terra firme forests (SC = 0.74, f1 = 63, f2 = 16). The spi-
der inventories were incomplete in all the sampled habitats, even con-
sidering the extrapolation of the number of individuals collected
(Figure 2a–c).
The curves of equal sampling effort indicate a wide overlap in the
species richness (q = 0) among the habitats (Figure 2a–c). Comparing
species richness at similar sample coverage (around 0.75) indicates
that islands had lower species richness compared with mainland habi-
tats (Figure 2a). Similar results were found when the alpha diversity is
measured as Shannon’s entropy (q = 1) or inverse of Simpson indices
(q = 2) using equal sampling effort (Figure 2a–c or sample coverage
(Figure 2d–f).
Distribution of individual among species was more equitable in
igap

o flooded forests (J = 0.94) and terra firme forests (J = 0.94) than in
fluvial islands (J = 0.87), indicating that spider assemblages in islands
present higher dominance levels than mainland habitats (KW = 10.23,

flooded forests comparison and
p = 0.02 for fluvial islands vs. igapo
p = 0.006 for fluvial islands vs. terra firme forests comparison).
Species composition was distinct between sampled habitats, a
pattern observed through relative abundance (Figure 3a) and presence

flooded forests
or absence data (Figure S1A). Fluvial islands and igapo
shared more species (Figures 3 and S1), which is expected given their
spatial and ecological proximity. Nevertheless, the species composi-
tion on river islands apparently is different from that found in the

flooded forests (Figures 3a and S1A).
neighbouring igapo
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6 DE CARVALHO ET AL.

TABLE 1 Results of analysis of similarity comparing species composition between studied habitats.

Dataset R global Paired comparisons R value

All species—abundance 0.76 IFF vs. TFF 0.61

IFF vs. FI 0.63
FI vs. TFF 0.95
Species (≥10 individuals)—abundance 0.74 IFF vs. TFF 0.63
IFF vs. FI 0.61
FI vs. TFF 0.92
All species—presence/absence 0.72 IFF vs. TFF 0.62
IFF vs. FI 0.57
FI vs. TFF 0.93
Species (≥10 individuals)—presence/absence 0.70 IFF vs. TFF 0.64
IFF vs. FI 0.55
FI vs. TFF 0.87

Note: R values close to 0 indicate great overlap in species composition, whereas values close to 1 indicate great distinction in species composition.

flooded forests; TFF, terra firme forest.
Abbreviations: FI, fluvial islands; IFF, igapo

TABLE 2 Indicator values of spider families identified as indicators of habitats in the Anavilhanas archipelago region, Central Amazon.

Families (individuals) Indicator of Fluvial islands
flooded forests

Igapo Terra firme forests

Salticidae (1640) FI 47.67 27.5 24.83

Araneidae (575) FI 42.37 28.71 28.92
Anyphaenidae (473) FI 42.79 37.65 9.356
Thomisidae (339) FI 53.91 9.485 30.53
Sparassidae (255) FI 42.86 30.92 16.47
Pisauridae (251) FI 72 9.634 0.4229
Pholcidae (129) FI 38.07 10.28 28.56
Scytodidae (124) IFF 19.41 44.93 9.764
Corinnidae (112) TFF 15.74 10.96 58.57
Tetragnatidae (61) FI 32.73 14.4 3.318
Theridiosomatidae (56) TFF 0.6406 5.848 68.57

Note: Bold numbers highlight significant ( p < 0.05) values in the permutation tests.

flooded forests; TFF, terra firme forest.
Abbreviations: FI, fluvial islands; IFF, igapo

The pattern of species composition differentiation between hab-
itats was confirmed in the analysis of similarity. The ANOSIM
R values were higher when the river islands were compared with the
terra firme forests, considering both abundance and presence or

flooded forests and fluvial
absence of data (Table 1). The igapo
islands, in contrast, tend to show greater overlap in species composi-
tion (Table 1). Despite this, the composition of spider species on the


fluvial islands was significantly different from the nearby igapo
flooded forests (Table 1).
The pattern of distinction in species composition between the
three habitats was little altered when the analyses were restricted to
the most-abundant species (Table 1). This indicates that the distinc-
tion in species composition between the habitats is not a sampling
artefact due to the inclusion of a high number of singletons in the
analysis. The distributions of the sites in the ordinations demonstrate
that spider species composition of fluvial islands remains distinct from

flooded forests and terra firme forests even when consider-
the igapo
ing only the most abundant species (Figures 3b and S1B).
Spider assemblages: Indicator taxa
Eleven spider families show significant association with a particular
habitat (Table 2). Spiders from Scytodidade family was an indicator of

flooded forests, whereas Corinnidae and Theridiosomatidae
igapo
were particularly abundant and frequent in terra firme forests
(Table 2). Another eight spider families were indicators of fluvial
islands (Table 2). Of the 23 species selected for the indicator species
analysis, 15 had significant associations with a particular habitat, most
of them to fluvial islands (Table 3). High IndVals were found for Scopo-
cira aff. carinata, Titidius aff. urucu, Cotinusa sp. 01 and Helvibis aff. ger-
maini (Table 3).
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SPIDERS IN Amazonian FLUVIAL ISLANDS 7

T A B L E 3 Indicator values of spider identified to species/morphospecies identified as indicators of habitats in the Anavilhanas archipelago
region, Central Amazon.

Species (individuals) Indicator of Fluvial islands
flooded forests

Igapo Terra firme forests

Scopocira aff. carinata (232) FI 81.86 2.56 0.12

Titidius aff. urucu (57) FI 62.05 2.62 0.00
Rishaschia amrishi (36) FI 27.83 5.50 0.00
Faiditus sp. 01 (29) FI 42.7 1.12 0.00
Achaearanea aff. hieroglyphica (25) FI 25.37 11.26 0.00
Thymoites sp. 01 (24) TFF 0.00 0.00 30
Cyclosa sp. 01 (24) FI 43.33 0.00 0.00
Cotinusa sp. 01 (21) IFF 0.45 35.87 0.00
Carapoia sp. 01 (20) FI 23.78 0.83 0.00
Helvibis aff. germaini (18) TFF 0.00 0.00 70
Lyssomanes amazonicus (18) FI 36.67 0.00 0.00
Castianeira sp. 01 (14) FI 20.8 1.03 1.73
Itata aff. tipuloides (13) IFF 0.52 27.89 1.18
Spintharus sp. 01 (12) IFF 1.26 53.77 0.00
Asaracus sp. 01 (11) FI 16.25 1.44 0.00

Note: Bold numbers highlight significant ( p < 0.05) values in the permutation tests.

flooded forests; TFF, terra firme forest.
Abbreviations: FI, fluvial islands; IFF, igapo

DISCUSSION
The number of spider species recorded in inventories throughout the
Amazon region is variable and dependent on sampling effort, methods
used and habitat types sampled, which precludes direct comparisons
between studies (Bonaldo et al. 2009a; Bonaldo et al. 2009b;
Höfer, 1990; Rego et al., 2009). However, a general comparison
between our results and previous findings can be useful to understand
the magnitude of the regional spider diversity in the Amazon.
The number of spider species recorded in Amazonian floodplain
habitats ranged from 130 to 384 species (Höfer, 1990; Rego
et al., 2009; Salvatierra, 2018). We recorded 223 species of spiders in

flooded forests), which is a
floodplain habitats (fluvial islands + igapo
level of diversity that is comparable to the aforementioned studies. In
contrast, spider fauna in well-studied sites of terra firme forests can
host 500–600 species (Bonaldo et al. 2009a; Bonaldo, Brescovit,
et al., 2009; Höfer & Brescovit, 2001). Our sampling revealed the
occurrence of only 104 species in terra firme forests, which indicates
that spider diversity in this habitat remains undersampled.
Our sampling was incomplete for all studied habitats. Indeed, to
obtain complete inventories for a group as diverse as spiders, it would
take years of sampling, application of complementary methods and
the collaboration of several specialists (Basset et al., 2012). Even with
this limitation, the patterns documented in our study were consistent,
since they were based on standardised samples and analytical
approaches that deal with incomplete samples (Chao et al., 2014;
Chao & Jost, 2012).
The typical attributes of island biotas have been identified in the
spider assemblages of the studied fluvial islands. Continuous habitats
must harbour a greater diversity of species than islands since they
occupy more extensive areas than islands (Rosenzweig, 1996; Whitta-
ker & Fernández-Palacios, 2009). Alpha diversity of spider on the flu-
vial islands was substantially lower than mainland habitats in all scales
of rarity represented by the Hill numbers.
A higher proportion of individuals of a few species on islands
compared with continuous habitats is also expected (Fattorini
et al., 2017; He & Legendre, 2002). Therefore, greater dominance
would be a consequence of a disproportionate increase in the abun-
dance of a few species that are well adapted to local ecological condi-
tions. Indeed, we observed lower equitability of individuals in spider
assemblages on river islands, which suggests that some species of spi-
ders may have been favoured by the insularity and have become dom-
inant on the islands. This trend is well exemplified by Scopocira aff.
carinata, the most abundant species (20% of samples) that was
recorded almost exclusively on the islands.
We expected greater compositional similarity between habitats
due to the proximity of the collection sites and the dispersive capacity
of the spiders. In particular, we anticipated a larger overlap in species
composition between river islands and igap

o flooded forests, which
are close and similar in vegetation structure and floristic composition
(Piedade et al., 2005). However, the species composition of the
islands was distinct enough to rule out the hypothesis that the island

flooded for-
spider fauna is only an impoverished subset of the igapo
ests and terra firme forests. This pattern of compositional distinction
could be an artefact caused by incomplete sampling and a large num-
ber of singletons. However, there is a clear compositional distinction
of the islands even considering only the most abundant species.
Differences in the spider assemblages between habitats were
identified even at the family level as had been reported by Privet et al.
(2018) and Lalagüe et al. (2022) in other environment types in French

8 DE
Guiana. For instance, spiders from the families Corinnidae and Theri-
diossomatidae were relatively more abundant in terra firme forests.
Most of the species in these families have highly sclerotized carapaces
and legs with few bristles (Bonaldo, 2000). In contrast, species of
Pisauridae and Anyphaenidae occurred with higher frequencies in flu-

flooded forests. Numerous spider species from
vial islands and igapo
these families are found in environments subject to flooding in micro-
habitats such as aquatic macrophytes (Bonaldo et al. 2009a;
Brescovit, 1996; Suter et al., 2003; von May et al., 2019). The natural
history and morphological traits of these spiders suggest that seasonal
flooding and insularity may select spiders with certain functional attri-
butes, thus favouring their adaptation to specific environments. This
aspect, however, is out of the scope of this study and deserves further
studies.
Some species of spiders were almost restricted to fluvial
islands, suggesting that some of them could be specialised in this
habitat type, as is reported for birds (Borges et al., 2019; Cintra
et al., 2007; Rosenberg, 1990). However, the analysis of habitat
specialisation must consider the limited understanding of the geo-
graphical distribution and habitat preferences of Amazonian spi-
ders. The species Lyssomanes amazonicus, for example, was
recorded only on islands (n = 18 individuals), but has a widespread
geographical distribution in the Brazilian Amazon, Bolivia and
Guyana (Bonaldo et al. 2009a; Carvalho & Gasnier, 2019;
Galiano, 1980). Therefore, the almost exclusive use of the islands
by some spiders could be a regional phenomenon for some species
of widespread distribution.
Genuine specialisation in the use of islands, however, should not
be discarded. For most species that have been associated with a par-
ticular habitat, there are other species of the same genus in another
habitat. These congeneric species that substitute each other between
habitats have similar morphology and likely occupy similar niches. For
example, the almost exclusive occurrence of Titidius aff. urucu on the
fluvial islands may be related to the presence of another three species
of the same genus in continuous habitats. Interspecific competition,
therefore, may be a mechanism underlying the possible specialisation
of some spider species in the use of fluvial islands, which also need
further investigation.
Implications for conservation
Events that alter the amplitude and duration of flooding regimes
potentially threaten species living in Amazonian riverine habitats
(Laranjeiras et al., 2021; Melo et al., 2021; Rosenberg, 1990). These
impacts could be even more relevant on fluvial islands due to their
spatial restriction and tiny occupied areas. If fluvial islands harbour
unique biological assemblages as suggested by our study and others
(Nunes et al., 2014; Rosenberg, 1990), this habitat deserves special
attention in conservation plans.
Amazonian floodplain habitats, including fluvial islands, are threat-
ened by human-driven activities with prominence to climate change
events and construction of hydroelectric dams (Castello et al., 2013).
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CARVALHO ET AL.
Recently, an increase in burning events in Amazonian floodplain for-
ests has been reported, which probably is linked to global climate
changes (Flores et al., 2017). Extreme flooding events have also been
recorded in the last decades in the Amazon basin (Espinoza
et al., 2022). Unfortunately, the responses of animal assemblages of
floodplains habitats to wildfires and extreme flooding are poorly
investigated (see Ritter et al., 2012).
Amazonian floodplains are also threatened by the intensification
in the construction of hydroelectric dams with strong ecological
impacts (Bueno et al., 2018; Cochrane et al., 2017; Forsberg
et al., 2017). There are 156 hydroelectric dams in operation, 21 under
construction and 277 planned in several tributaries of the Amazon
basin (Macedo & Castello, 2015). Hydroelectric dams alter the flood-
ing dynamics in the vicinity of the reservoirs, thus resulting in the loss
of fluvial islands and other floodplain habitats that will end up perma-
nently underwater. Loss of fluvial islands could affect regional popula-
tion dynamics of species specialised in these habitats, an aspect that
also deserves further investigation.
Fluvial islands could be affected by hydroelectric dams even if
they are located inside the limits of protected areas. For example, the
Brazilian government is planning to construct a dam in the Branco
River, the largest tributary of Negro River located a 100 km upriver
from our study sites (Laranjeiras et al., 2021). Damming Branco River
channel will drastically reduce the sediment input that maintains the
fluvial islands localised in the protected areas of the lower Negro
River (Laranjeiras et al., 2021), including our study sites in the Anavil-
hanas National Park.
CONC LU SION
Patterns shared by island biotas around the world were first docu-
mented in spider assemblages that inhabit river islands in the Ama-
zon. The taxonomic alpha diversity of spiders was lower on islands
compared with mainland habitats. The spider assemblages of the
islands also showed a higher dominance of a restricted set of species.
Fluvial islands effectively contribute to regional diversity since the
composition of the spider species on the islands was distinct from
the continuous mainland habitats. Habitat specialisation may be
involved in the differentiation of island spider assemblages, which is
an aspect that requires further studies. Events that alter the flood
pulse in Amazonian rivers could affect the processes
(e.g., sedimentation) that create and maintain floodplain habitats
such as river islands, which, in turn, could threaten their biologically
distinctive spider assemblages.
AUTHOR CONTRIBU TIONS
Thiago Gomes de Carvalho: Conceptualization; methodology; formal
analysis; data curation; writing – review and editing. Thierry Ray Gasnier:
Conceptualization; methodology; writing – review and editing; data cura-
tion; supervision. Sérgio Henrique Borges: Funding acquisition;
writing – original draft; writing – review and editing; formal analysis; pro-
ject administration; resources; supervision.

SPIDERS IN Amazonian FLUVIAL ISLANDS
ACKNOWLEDGEMEN TS
This manuscript results from the master’s dissertation developed by

s-graduação em
Thiago Gomes de Carvalho in the Programa de Po
Zoologia at the Universidade Federal do Amazonas (UFAM), which is
supported by Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES). This study was financed in part by the Coordenação
de Aperfeiçoamento de Pessoal de Nível Superior–Brasil (CAPES)–
Finance Code 001. During our fieldwork in Anavilhanas National Park,
we had logistical support provided by the technical team from the
Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio),
especially by Priscila Santos, Enrique Salazar, Paula Pinheiro,
Adenilson dos Santos and Maria Izabel da Silva. We are also grateful
to Gerley de Vasconcelos Anicacio, Ralison Vasconcelos Anicacio and
Phillip Klauvin for their enthusiastic field support during the spider
sampling. Eduardo Martins Venticinque, Fabrício Baccaro and Lidianne
Salvatierra made valuable suggestions regarding our project. During
the development of this study, Thiago Gomes de Carvalho received a
grant from Fundação de Amparo à Pesquisa do Estado do Amazonas
(FAPEAM) and Sérgio Henrique Borges was supported by FAPEAM
through grants from Programa Fixam (call 017/2014) and Programa
FAPEAM Produtividade em CT&I (call 013/2022). Financial support
for fieldwork was provided by FAPEAM through the Universal
Amazonas (call 002-2018) and Programa Fixam (call 017/2014).
FUND ING INFORMATION
This work was supported by Fundação de Amparo à Pesquisa do
Estado do Amazonas (FAPEAM) through Programa Fixam (call
017/2014) and Universal Amazonas call (call 002-2018).
CONF LICT OF IN TE RE ST ST AT E MENT
The authors declare no conflicts of interest.
DATA AVAI LAB ILITY S TATEMENT
The data that supports the findings of this study are available in the
supplementary material of this article.
ORCID
Sérgio Henrique Borges https://orcid.org/0000-0002-1952-4467
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SPIDERS IN Amazonian FLUVIAL ISLANDS 11

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