Neotropical Ichthyology Original article [Link]

1590/1982-0224-2023-0135

Spatial and trophic structure drive

ecomorphological patterns of fish
assemblages in oligotrophic headwater
streams of northwestern Amazonia
Correspondence:
Francisco Javier Luque
fjluquem@[Link] Francisco Javier Luque1 and Gabriel J. Colorado Z.2

This study evaluated the ecomorphological patterns of fish assemblages in

oligotrophic headwater streams of northwestern Amazonia. Our aims were: 1)
identify correlations between morphological variables of fish species and habitat
use, trophic structure and distance to the main river; 2) examine ecomorphological
patterns that reflect ecological aspects of locomotion, habitat use and trophic
ecology. A total of 34 species were analyzed using 27 ecomorphological indices
to test if they were related to fish assemblage. Significant correlations were
found between anatomical traits and trophic guilds and habitat use regardless of
phylogenetic relationships. Two trends were identified in the ecomorphological
space. First, nektonic insectivores-piscivores with lateral compressed bodies,
large eyes, superior-terminal mouth without barbels and larger anal fins were
Submitted December 13, 2023 associated to species with continuous swimming, and the use of vision to capture
Accepted March 26, 2025 prey. Second, benthic periphytivores-invertivores with depressed bodies, small
Epub June 6, 2025 eyes, ventral or inferior mouth with presence of barbels, and developed pectoral
fins were related to species that exploit streams substrate. This study shows that
Associate Editor Caroline Arantes fish assemblages in the oligotrophic headwater streams are ecomorphologically
Section Editor Fernando M. Pelicice structured by trophic guilds and habitat use. These findings highlight the use of
Editor-in-chief José Birindelli ecomorphology as a tool to predict ecological patterns in these streams.

Keywords: Guaviare, Habitat use, Morphology, Phylogeny, Trophic guilds.

Online version ISSN 1982-0224

Print version ISSN 1679-6225

1 Grupo en Ecología y Conservación de Fauna y Flora Silvestre, Departamento de Biología, Universidad Nacional de Colombia.
Carrera 45 # 26 – 85, 111321, Bogotá D.C, Colombia. fjluquem@[Link] (corresponding author).
Neotrop. Ichthyol.
2 Grupo en Ecología y Conservación de Fauna y Flora Silvestre, Departamento de Ciencias Forestales, Universidad Nacional de
vol. 23, no. 2, Maringá 2025 Colombia. Carrera 65 # 59a – 110, Medellín, Colombia. gjcoloradoz@[Link].

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 Fish ecomorphology in oligotrophic streams

Este estudio evaluó los patrones ecomorfológicos de los ensamblajes de peces

en los ríos oligotróficos de cabecera al noroeste de la Amazonía colombiana.
Nuestros objetivos fueron: 1) identificar correlaciones entre las variables
morfológicas de los peces y el uso de hábitat, estructura trófica y distancia al río
principal; 2) examinar patrones ecomorfológicos que podrían reflejar aspectos
ecológicos de la locomoción, uso de hábitat y ecología trófica. Un total de 34
especies fueron analizadas usando 27 índices ecomorfológicos para probar si estos
estaban relacionados con el ensamblaje de peces. Se encontraron correlaciones
significativas entre rasgos anatómicos y gremios tróficos y uso de hábitat
controlando las relaciones filogenéticas. Dos tendencias fueron identificadas
dentro del espacio ecomorfológico. Primera, nectónicos insectívoros-piscívoros
con cuerpos lateralmente comprimidos, ojos grandes, boca terminal-superior
sin barbicelos y aletas anales grandes asociados con especies de nado continuo y
uso de visión para capturar presas. Segunda, bénticos perifitovoros-invertívoros
con cuerpos deprimidos, ojos pequeños, boca ventral o inferior con presencia de
barbicelos y aletas pectorales desarrolladas asociadas con especies que explotan el
sustrato de los ríos. Este estudio evidencia que los ensamblajes de peces en los ríos
oligotróficos de cabecera se encuentran estructurados por los gremios tróficos y el
uso de hábitat. Estos resultados resaltan la importancia del uso de la ecomorfología
como una herramienta para predecir patrones ecológicos en estos ríos.

Palabras clave: Filogenia, Gremios tróficos, Guaviare, Morfología, Uso de hábitat.

INTRODUCTION

Ecomorphology analyzes the relationships between morphological traits, ecology,

and how they relate to the environment (Bock, 1994; Wainwright, Richard, 1995).
The main premise of ecomorphology suggests that species with similar morphology
share similar resources (Fernández et al., 2012). Under this conceptual basis, organismal
morphology allows the identification of biological roles, the environment where the
species live in (Wootton, 1990), how species exploit and uses resources (Wainwright,
Richard, 1995; Hugueny, Pouilly, 1999), the niche they occupy and, in general, how
species assemblage is structured (Conde-Saldaña et al., 2017) through species anatomy
(Gatz Jr., 1979). Thus, ecomorphological analysis provides a framework to understand
and predict ecological patterns for the use of spatial and trophic resources.
Nevertheless, several studies have shown a lack of consensus on whether morphology
is correlated to ecology (Bhat, 2005; Oliveira et al., 2010). This disagreement has
extended to the paraphyletic group of fishes, the most taxonomically diverse vertebrate
group on earth, for which ecomorphology represents a cornerstone discipline of
research given their variety of forms and functions. For example, evidence suggests
that depressed bodies exploit benthonic habitats, and that compressed bodies inhabit
structured habitats (Gatz Jr., 1979; Cochran-Biederman, Winemiller, 2010; Soares et al.,
2013) or that long digestive tracts are strongly related to herbivory and, in turn, head
and mouth size is related to prey size (Ferreira, 2007; Cochran-Biederman, Winemiller,

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Francisco J. Luque and Gabriel J. Colorado Z.

2010; Oliveira et al., 2010). Nevertheless, other studies differed in these findings by
suggesting a weak relationship between the morphology of the organism and its
environment (Motta et al., 1995; Casatti, Castro, 2006; Conde-Saldaña et al., 2017).
This discrepancy has been associated with phylogenetic influence (Douglas, Matthews,
1992; Motta et al., 1995), diverse statistical approaches to test hypothesis (Ricklefs, Miles,
1994), varied and inadequate selection of anatomical traits reliable to predict diet and
habitat (Oliveira et al., 2010), and the influence of environmental and habitat variables
(Lombarte et al., 2012; Oikonomou et al., 2018; Luiz et al., 2022).
From headwaters to downstream, streams increase in size, discharge, number of habitats
and a myriad of physical, chemical, and ecological parameters along the longitudinal
gradient within the dendritic stream network (Bistoni, Hued, 2002; Grenouillet et al.,
2004). Some studies have investigated how fish assemblages and ecomorphological
patterns respond to altitudinal changes (Conde-Saldaña et al., 2017), current velocity
(Casatti, Castro, 2006), hydrological gradient (Luiz et al., 2022), and habitat features
(Oliveira et al., 2010; Prado et al., 2016; Oikonomou et al., 2018). However, the effect
of distance from source on ecomorphological patterns have received little attention,
some studies have found that richness, trophic guild proportions and fish assemblages
changed along the position of tributaries along the longitudinal continuum of tropical
and temperate streams (Rahel, Hubert, 1991; Oberdorff et al., 1993; Fialho et al., 2007).
Indeed, distance to large rivers has shown to be relevant in explaining the diversity and
structure of fish assemblages at different spatial scales (e.g., Stegmann et al., 2019). For
example, some adventitious streams may have richer fish communities than similar-size
streams located further up in the network due to migration from larger rivers (Schaefer,
Kerfoot, 2004).
Despite the vast diversity and endemism of fishes in tropical freshwater ecosystems,
most ecomorphological studies have been carried out on temperate freshwater
ecosystems (Bhat, 2005). However, some ecomorphological research have aimed to
describe these types of relationships in different Amazonian aquatic environments such
as white waters and its floodplains ( Pouilly et al., 2003; Ibañez et al., 2007; Mérona
et al., 2008), but a few had addressed such studies on oligotrophic headwater streams
(Ramírez et al., 2015). These are small, nutrient poor, dark water acidic streams that
support a diverse and rich fish fauna, highly dependable on allochthonous inputs and
abiotic conditions such as heavy rains that dramatically change water levels (Correa,
Winemiller, 2018). Therefore, oligotrophic headwater streams represent an opportunity
to test ecomorphological hypotheses encompassing an understudied geographical area.
In this study we examined ecomorphological patterns of fish assemblages in
oligotrophic streams. Our aims were to: 1) Identify and assess correlations between
morphological variables of fish species with habitat use, trophic guilds, and distance to
the Guaviare River while controlling for phylogenetic relationships, and 2) describe
possible ecomorphological patterns that reflect ecological aspects of locomotion, habitat
use, and trophic ecology. We hypothesized that species with similar morphology will
share both habitat and trophic guild, as well as exhibit similar functional roles, feeding,
and locomotion behaviors. Additionally, fish assemblages close to the Guaviare River
will present higher phylogenetic and functional richness than those closer to headwater
(Stegmann et al., 2019). In this sense, the morphology of the species would differ
among trophic guild by resource use, whereas it would differ in habitat use by physical,
chemical, and environmental characteristics (Oliveira et al., 2010).

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 Fish ecomorphology in oligotrophic streams

MATERIAL AND METHODS

Study area. This study was conducted in headwater oligotrophic streams in the Guaviare
River basin near the municipality of San José del Guaviare, Guaviare Department,
northwestern part of the Colombian Amazon. The sampling area is located within the
Guiana shield region, corresponding to a transition zone between the Amazon rain
forest and the Orinoco grasslands (Huber, 2006; Medina-Rangel et al., 2019). The area
has a tropical rainforest regime, with an annual precipitation average of 2,800 mm and
a monomodal regime (dry period from November to February). Average temperature
and humidity are 25.7 oC and 78–90%, correspondingly (Cárdenas-López et al., 2008).
Streams headwaters are located within the Serranía de la Lindosa, a protected area with
an extension of 12,000 ha. According to Lasso (2014), these streams are part of a system
of white waters, affluents to the Guaviare River and part of the Orinoco River basin.
Vegetation is composed by riparian forests, rocky outcrops shrubs and forests, white
sand stands, and grasslands (Atuesta-Ibargüen, 2019).

Field sampling. 15 sampling stations were established from ~1 to 30 km (mean ± SD:

14.53 km ± 7.60) from the main stream (Guaviare River), ranging from 185 to 285 m in
elevation (Tab. 1). Station selection was based on the following criteria: 1) landowner
access permission; 2) physical security; and 3) good accessibility to ensure repeatability.
At each station, a reach of 350–400 m was established. Minimum distance between
reaches was 3 km, maximizing independence among study reaches. Each reach was
divided into three sections (i.e., Upstream, Midstream, and Downstream) following the
methods used by (Torres-Bejarano et al., 2022) (Fig. 1).

TABLE 1 | Physical characteristics in 15 sampling stations in oligotrophic headwater streams of northwestern Amazonia, Guaviare Department,
Colombia.

Distance Mean Geographical

Sampling Elevation Mean Mean Mean flow
from river Habitat type flow velocity coordinates
Station (m) depth (m) width (m) (m³/s)
(km) (m/s)
N W
Yam_Med 23.17 235 Riffle 0.50 ± 0.18 0.31 ± 0.16 5.58 ± 0.79 0.8649 2°27'13.70"S 72°45'17.80"W
Yam_Int 29.68 281 Riffle 0.36 ± 0.04 0.44 ± 0.05 5.37 ± 1.45 0.850608 2°27'25.90"S 72°42'40.30"W
Yam_Con 1.17 200 Pool 0.80 ± 0.64 0.06 ± 0.61 11.58 ± 4.83 0.55584 2°35'26.70"S 72°50'34.20"W
Ret_Med 20.95 219 Pool 0.60 ± 0.18 0.05 ± 0.07 9.48 ± 0.54 0.2844 2°30'29.00"S 72°43'5.00"W
Ret_Int 17.98 242 Riffle 0.48 ± 0.26 0.33 ± 0.18 5.34 ± 4.72 0.845856 2°31'32.70"S 72°42'48.80"W
Ret_Con 14.4 212 Pool 0.60 ± 0.32 0.11 ± 0.15 5.51 ± 2.28 0.36366 2°33'13.10"S 72°42'44.90"W
Lin_Med 19.19 219 Pool 0.83 ± 1.11 0.08 ± 0.16 7.45 ± 3.46 0.49468 2°29'21.71"S 72°39'37.92"W
Lin_Int 12.65 203 Pool 0.96 ± 0.08 0.02 ± 0.17 20.57 ± 2.22 0.394944 2°30'26.40"S 72°38'27.10"W
Lin_Con 16.65 208 Riffle 0.40 ± 0.02 0.17 ± 0.1 14.96 ± 6.88 1.01728 2°29'45.60"S 72°39'2.20"W
Agu_Med 2.43 189 Pool 1.07 ± 0.46 0.02 ± 0.07 23.48 ± 2.42 0.502472 2°31'25.80"S 72°37'7.50"W
Agu_Int 7.23 185 Pool 0.73 ± 1.03 0.03 ± 0.54 14.46 ± 12.2 0.316674 2°34'45.50"S 72°36'4.90"W
Agu_Con 11.04 200 Pool 1.08 ± 0.17 0.03 ± 0.11 8.86 ± 1.83 0.287064 2°32'55.00"S 72°36'57.30"W
Cri_Med 17.94 196 Riffle 0.26 ± 0.09 0.38 ± 0.11 2.1 ± 0.85 0.20748 2°32'47.80"S 72°45'10.70"W
Cri_Int 13.27 203 Riffle 0.43 ± 0.30 0.5 ± 0.13 2.94 ± 0.71 0.6321 2°32'58.10"S 72°44'21.70"W
Cri_Con 10.27 200 Riffle 0.54 ± 0.36 0.26 ± 0.08 3.97 ± 0.98 0.557388 2°33'15.50"S 72°43'56.60"W

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Francisco J. Luque and Gabriel J. Colorado Z.

FIGURE 1 | Study area with sampling station location in oligotrophic headwater streams of northwestern Amazonia, Guaviare Department,
Colombia.

Environmental variables. Channel width was considered as the distance between

stream margins and measured with a measuring tape. Stream velocity was calculated
using a FlowWatch (JDC electronic SA, Switzerland) and stream flow by multiplying
mean width (m), mean depth (m), and mean velocity (m/s). Two habitats were evaluated,
riffles and pools. Riffles were characterized by fast moving water and shallower depth,
whereas pools showed slower flows and deeper waters (Tab. 2).

Fish collection and morphological data. Fish were collected at study reaches once
during the wet and once during the dry season using active seining (30 m x 2 m with 2.5
cm mesh) from downstream to upstream and passive gill netting (15 m x 3 m with 2.5
cm mesh) at both ends of the reach closing it and preventing the entry and exit of fish.
Our protocols followed those proposed by Anjos, Zuanon (2007) and Torres-Bejarano
et al. (2022), which support that ~240 m is sufficient to adequately represent fish richness
in small Amazonian streams. Our sampling efforts included 70–90 min collections over
~300 m2 at each study reach. The fish collected were anesthetized and euthanized with

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 Fish ecomorphology in oligotrophic streams

TABLE 2 | Physical characteristics of the evaluated habitats.

Habitats Characteristics

Mean depth (m) Mean width (m) Mean flow (m3/s) Mean velocity (m/s)

Riffles 0.44 4.78 0.7 0.34

Pools 0.83 8.85 0.39 0.04

MS–222 (200 mg/L), and frozen for posterior analysis in the laboratory. All individuals
were identified to species using taxonomic keys (Géry, 1977; Gálvis et al., 2006, 2007;
Kullander, 2006; Urbano-Bonilla et al., 2017), and expert assistance when necessary.
Species were classified into trophic guilds (i.e., detritivores, insectivores, invertivores,
periphytivores, and piscivores) using published information (Gálvis et al., 2006, 2007;
van der Sleen, Albert, 2018; Froese, Pauly, 2023).
Twenty three linear morphometric measurements and six areas were taken on the left
side of the individuals using a digital caliper (0.01 mm precision), whereas areas of eyes
and fins were obtained from photographs, and calculated using AutoCAD software (Fig.
2). Only species with more than five adult individuals were measured, considering that
rare species could reflect non-interactive components of main community (Winemiller,
1991), small samples limit the detection of specific morphological variations (Oliveira
et al., 2010), and ontogenetic growth can promote significant changes in body shape
(Bower, Piller, 2015), habitat use (Sagnes et al., 1997) and feeding habits (Oliveira et al.,
2010; Prado et al., 2016). After all measurements were taken, specimens were fixed with
10% concentration formalin and then transferred to 70% ethanol solution and deposited
in the Ichthyological collection of Instituto de Ciencias Naturales (ICN) of Universidad
Nacional de Colombia, Bogotá, Colombia.
Based on these criteria, 34 of 62 species captured were measured and considered
for further analysis (Tab. 3). Linear measurements and areas were used to calculate
27 ecomorphological indices related to locomotion, habitat use, and trophic ecology
(Tab. 4). Additionally, two ecological traits that do not involve measurements were
considered for the analysis: presence of barbels and mouth position. Both traits are
related to trophic guild and habitat use (Tab. 5). These ecomorphological traits are
unitless ratios, so they are not correlated with body size (Oikonomou et al., 2018),
allowing comparisons between species or individuals of different sizes and focus on
function and ecological aspects of morphology while minimizing the confounding
effects of body size (Bower, Piller, 2015) .

Data analyses. Ecomorphological structure. To test the hypothesis that species

with similar morphology will share both habitat and trophic guild, independent of
phylogeny (taxonomic), a Mantel test analysis was performed. This test assesses the
correlation between two distance matrices and the statistical significance is obtained
by n randomizations of the input distance matrices. Thus, the analysis seeks to test
the hypothesis that the matrices are not linearly or monotonically related (Legendre,
Legendre, 2012). To conduct this analysis, we generated a distance matrix containing
the morphological information of the fish assemblage (i.e., morphological matrix) and

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Francisco J. Luque and Gabriel J. Colorado Z.

FIGURE 2 | Linear morphometric measurements and calculated areas: standard length (SL), dorsal-
fin length (DL), caudal peduncle length (Lcp), caudal fin length (CL), pectoral-fin length (PTL), pelvic-
fin length (PvL), anal-fin length (AL), mouth width (MW), maximum body height (HMB), midbody
height (BHM), dorsal fin height (DH), caudal peduncle depth (Dcp), caudal fin height (CH), pectoral fin
height (Pth), pelvic fin height (PVH), anal fin height (AH), head height (HH), eye height (EH), mouth
width (MW), mouth height (MH), maximum head width (MHW), maximum body width (MBW), caudal
peduncle width (WCP), eye area (EA), dorsal fin area (Dfa), caudal fin area (CA), anal fin area (Afa), pelvic
fin area (PeLA), and pectoral fin area (Pca). Based on a specimen of Astyanax bimaculatus.

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 Fish ecomorphology in oligotrophic streams

TABLE 3 | Fish species of the oligotrophic headwater streams in northwestern Amazonia with trophic guilds, habitat association and distance
from source group.

Distance
Measured Habitat Trophic
Order/Family/Species Code to river
specimens type guild
(km)
Characiformes

Acestrorhynchidae

Acestrorhynchus falcatus (Bloch, 1794) Ac_fal 5 Pool 0-5 Piscivore

Bryconidae

Brycon whitei Myers & Weitzman, 1960 Br_whi 11 Pool 6 - 10 Piscivore

Acestrorhamphidae

Astyanax integer Myers, 1930 As_int 20 Pool >15 Insectivore

Astyanax bimaculatus (Linnaeus, 1758) As_bim 20 Pool 0-5 Omnivore

Astyanax sp.2 A_sp2 20 Pool 6 - 10 Insectivore

Bario oligolepis (Günther, 1864) Mo_oli 17 Pool 6 - 10 Insectivore

Hemigrammus barrigonae Eigenmann & Henn, 1914 He_bar 20 Riffle >15 Omnivore

Hyphessobrycon acaciae García-Alzate, Román-Valencia & Prada-Pedreros, 2010 Hy_aca 20 Riffle 0-5 Omnivore

Hyphessobrycon taguae García-Alzate, Román-Valencia & Taphorn, 2010 Hy_tag 20 Riffle 11 - 15 Omnivore

Characidae

Charax metae Eigenmann, 1922 Ch_met 10 Pool 11 - 15 Piscivore

Stevardiidae

Creagrutus calai Vari & harold, 2001 Cr_cal 20 Riffle >15 Insectivore

Moenkhausia comma Eigenmann, 1908 Mo_com 15 Pool 0-5 Insectivore

Moenkhausia mikia Marinho & Langeani, 2010 Mo_mik 16 Pool 0-5 Insectivore

Tyttocharax metae Román-Valencia, García-Alzate, Ruiz-C. & Taphorn, 2012 Ty_met 5 Riffle 6 -10 Omnivore

Curimatidae

Cyphocharax oenas Vari, 1992 Cy_one 5 Pool 6 - 10 Detritivore

Steindachnerina argentea (Gill, 1858) St_arg 6 Pool 0-5 Detritivore

Erythrinidae

Hoplerythrinus unitaeniatus (Spix & Agassiz, 1829) Ho_uni 5 Pool 6 - 10 Piscivore

Hoplias malabaricus (Bloch, 1794) Ho_mal 8 Pool 6 - 10 Piscivore

Iguanodectidae

Bryconops giacopinii (Fernández-Yépez, 1950) Br_gia 20 Pool 11 - 15 Insectivore

Lebiasinidae

Copella arnoldi (Regan, 1912) Co_arn 20 Pool 11 - 15 Omnivore

Pyrrhulina lugubris Eigenmann, 1922 Py_lug 20 Pool 11 - 15 Insectivore

Cichliformes

Cichlidae

Aequidens tetramerus (Heckel, 1840) Ae_tet 18 Pool 0-5 Invertivore

Apistogramma hoignei Meinken, 1965 Ap_hoi 16 Pool 11 - 15 Invertivore

Bujurquina mariae (Eigenmann, 1922) Bu_mar 15 Pool 0-5 Omnivore

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Francisco J. Luque and Gabriel J. Colorado Z.

TABLE 3 | (Continued)

Distance
Measured Habitat Trophic
Order/Family/Species Code to river
specimens type guild
(km)
Satanoperca mapiritensis (Fernández-Yépez, 950) Sa_map 7 Pool 0-5 Invertivore

Saxatilia alta (Eigenmann, 1912) Cr_alt 8 Pool 6 - 10 Piscivore

Cyprinodontiformes

Rivulidae

Anoblepsoides sp. An_sp1. 19 Pool 6 - 10 Omnivore

Siluriformes

Auchenipteridae

Duringlanis romani (Mees, 1988) Ce_rom 7 Riffle 0-5 Invertivore

Callichthyidae

Hoplisoma melini (Lönnberg & Rendhal, 1930) Co_mel 6 Riffle 0-5 Invertivore

Loricariidae

Ancistrus triradiatus Eigenmann, 1918 An_tri 6 Riffle >15 Detritivore

Farlowella vittata Myers, 1942 Fa_vit 17 Riffle 0-5 Detritivore

Rineloricaria eigenmanni (Pellegrin, 1908) Ri_eig 18 Riffle 0-5 Detritivore

Heptapteridae

Pimelodella metae Eigenmann, 1917 Pi_met 15 Pool 0-5 Invertivore

Trichomycteridae

Ituglanis metae (Eigenmann, 1917) It_met 17 Riffle 0-5 Insectivore

compared it against a set of generated matrices comprising phylogenetic, habitat use,

trophic guilds, and distance to river information, as follows:
Morphological matrix was calculated using a Euclidean distance matrix made for
each pair of species and based on their ecomorphological indices. This matrix was
generated using the average of each species and the decostand function of Vegan
package on Rstudio software (R Development Core Team, 2011).
Phylogenetic matrix was constructed following Douglas, Matthews (1992) and
Conde-Saldaña et al. (2017). A value of one was attributed to the pairs of congeneric
species; a value of 1.5 to those in different genera but same subfamily; a value of two to
those in different subfamily but same family; a value of three to those in different families
but in the same order; a value of 4 to those of different order but same superorder; and
a value of 5 in for those of different superorder. We used this approach as a surrogate
of phylogenetic relationships since a complete phylogenetic tree of Colombian amazon
freshwater fish species is not currently available.
Habitat use matrix was constructed following Conde-Saldaña et al. (2017), using
abundance as a predictor of relationship between fish and their habitats (Rosenfeld,
2003). Since abundance did not differ between dry and wet seasons (non-parametric
Kruskall-Wallis, H (1) = 0.0348; p = 0.851), the data was pooled to establish the species
association in each habitat type (riffle and pools). The indicator value method (IndVal)
Dufrene, Legendre (1997) was used to establish species habitat association; thus, each
species was assigned to a type of habitat according to the results of the maximum

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 Fish ecomorphology in oligotrophic streams

TABLE 4 | Ecomorphological indices used in the present study and their respective code, formulae, ecological relation, and biological
interpretation.

Ecomorphological index Code Code Formula Related to Biological interpretation

High values indicates that fish is laterally compressed, which could be
Compression index Com_ind Ci MBH/MBW Habitat use related to the exploit of slow water velocity habitats (Watson, Balon,
19984)
Low values indicates that fish presents a depressed body, which could be
Depression index De_ind Di BMH/MBH Habitat use
related to the exploit of bottom water column (Watson, Balon, 1984)
Low values could indicate fish occupy high velocity habitats (Gatz Jr.,
Relative depth Re_dep Rd MBH/SL Habitat use
1979; Watson, Balon, 1984)
High values indicate dorsally positioned eyes, often found on benthic
Eye position Eye_pos EP EH/Hd Habitat use species. Low values indicates laterally positioned eyes, found on nektonic
species (Poully et al., 2003; Watson, Balon, 1984)
Re_area_ High values are related to benthic species whereas lower values to
Relative area of pelvic fin Rap PeL/(SL)2 Habitat use
pfin nektonic species (Breda, 2005; Gatz Jr., 1979)
High values are related to nektonic habits whereas lower values to
As_ra_
Aspect ratio of pelvic fin Arp (PeL)2/PeA Habitat use benthic habits since fins are use as support on the substrate (Breda, 2005;
pfin
Gatz Jr., 1979)
High values are related to caudal propulsion efficiency for fast
Caudal peduncle trottle Ca_pen_ro Cpt Cfd/Cpd Locomotion
movements through a reduction in drag (Luiz et al., 2022)
High values are present in species have active and continuous swimming
Aspect ratio caudal fin As_ra_cfin Arc Cfd/Cfa Locomotion
habits (Balon et al., 1986)
Re_area_ Larger values are associated with active and continuous swimming
Relative area caudal fin Rac Cfa/(SL)2 Locomotion
cfin habits (Balon et al., 1986)
Caudal peduncle ca_pen_ High values are present in species with low swimming activity and low
Cpc Locomotion
compression index comin manuverability (Gatz Jr., 1979; Watson, Balon, 1984)
Larger values are related to species with good swimming capacity and
Relative length caudal Re_len_
Rlc Lcp/SL Locomotion fish that inhabit fast water flow habitats (Breda , 2005; Watson, Balon,
peduncle capend
1984; Winemiller, 1991)
Relative height caudal Rehig_ Low values are associated to species withhigh manuvelavility capacity
Rhc Hcp/MBH Locomotion
peduncle capend (Oliveira et al., 2010; Winemiller, 1991)
Relative width caudal Re_wid_ Low values areassociated to species with less capacity for continuous
Rwc Wcp/MBW Locomotion
peduncle capend swimming (Winemiller, 1991)
Re_area_
Relative area dorsal fin Rad Dfa/(SL)2 Locomotion High values provide greater stabilization (Breda, 2005; Gosline, 1971)
dfin
Re_area_ High values provide greatar stabilization and movement capacity (Breda,
Relative area anal fin Raa Afa/(SL)2 Locomotion
afin 2005; Wolff, 2008)
High values are related to the capacity to perfrm fast forward and
Aspect ratio anal fin As_ra_afin Ara Afl/Afa Locomotion
backwards movement (Breda, 2005; Wolff, 2008)
Low values are present in species that inhabit slow flowing waters.
Re_area_ Locomotion / High values are present in slow swiming species or species with benthic
Relative area pectoral fin Rape Pca/(SL)2
pecfin habitat use habitats that deflect water to stay as close as possible to the substrate
(Watson, Balon, 1984)
As_ra_ High values are present in species that swim continuously, can reach
Aspect ratio pectoral fin Arpe (Pcl)2/Pca Locomotion
pecfin higher speeds or migrate (Breda, 2005; Watson, Balon, 1984)
Use as an aprroximation for vision capacity. Larger eyes are related to
Relative eye size Rel_eye_si Res Ea/(SL)2 Trophic
food detection and its use for predatory activity (Poully et al., 2003)
Re_ Related to predatory capacity. Larger values are related to species that
Relative height of head Rhh MHH/MBH Trophic
hehead feed on larger preys (Wolff, 2008)
Re_ Related to predatory capacity. Larger values are related to species that
Relative width of head Rwh MHW/MBW Trophic
wihead feed on larger preys (Soares et al., 2013)
Related to predatory capacity, indicating the size of prey items. High
Relative height of mouth Re_hemou Rhm MMH/MBH Trophic
values are indicate species that feed on larger preys (Watson, Balon, 1984)
Re_ Related to predatory capacity, indicating the size of prey items. High
Relative width of mouth Rwm MWM/MBW Trophic
wimou values are indicate species that feed on larger preys (Watson, Balon, 1984)
(Mmw*Mmh)/ Related to predatory capacity, indicating the size of prey items. High
Oral gap size O_gap_si Ogs Trophic
(SL)2 values are indicate species that feed on larger preys (Luiz et al., 2022)
Related to predatory capacity, indicating the size of prey items. High
Oral gap shape O_gap_sh Osgh Mmh/Mmw Trophic
values are indicate species that feed on larger preys (Luiz et al., 2022)
Presence of barbels are related to species that primarly feed and inhabit
0 = ausent Trophic /
Barbels Barb Bar the bottom of the water column (Hugueny, Poully, 1999; Conde-Saldaña et
1 = present habitat use
al., 2017)
1 = superior
Position of mouth indicates the location in the water column where the
2 = terminal
Trophic / species obtain its food. Inferior and vental mouths are present in benthic
Mouth position Mo_pos Mp 3 = subterminal
habitat use species where as terminal and superior mouth are present in nektonic
4 = inferior
species (Gatz Jr., 1979; Conde-Saldaña et al., 2017)
5 = ventral

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Francisco J. Luque and Gabriel J. Colorado Z.

indicator value for each type of habitat (maxcls). This analysis was performed using
the IndVal function of the labdsv package of Rstudio (R Development Team, 2011). A
value of one was given to the pair of species that belong to the same habitat type and a
value of two for species that did not fulfill this condition (Soares et al., 2013).
Trophic guild matrix was generated using the trophic guild assign to each species; a
value of one was attributed to the pair of species belonging to the same guild and a value
of two for species belonging to different guilds (Soares et al., 2013).
Distance to river matrix was constructed following Conde-Saldaña et al. (2017). Four
groups with different distances to river were generated by taking the square root of the
number of sampling locations (Tab. 2). Abundance data in each sampling site was used
to perform the IndVal analysis and classification in each distance to river group was
done using the maxcls option from the IndVal function of the labdsv package of Rstudio
(R Development Team, 2011). The matrix was constructed giving a value of one to
species belonging to the same distance to river group whereas a value of two was given
to the pairs of species that did not satisfy this condition (Soares et al., 2013).
To avoid false correlations between morphology and trophic guild, habitat use, and
distance to river structures due to phylogenetic effects (Winemiller, 1991; Douglas,
Matthews, 1992; Neves et al., 2015), we assessed partial correlations between two matrices
(i.e., morphological vs. habitat use, morphological vs. trophic guilds, and morphological
vs. distance to river) while controlling for the effect of a third matrix (phylogenetic) using
a partial Mantel test. Mantel and partial Mantel tests were carried out using mantel and
[Link] functions of vegan package Rstudio (R Develompent Core Team, 2011).

Ecomorphological patterns. In order to examine and identify patterns of fish

assemblages distribution in the multivariate ecomorphological space and seek for
interspecific differences in ecomorphological traits (i.e., locomotion, habitat use, and
trophic ecology), we used a Principal Components Analysis (PCA). PCA was applied on
the correlation matrix composed by the mean values of the 27 ecomorphological indices
previously transformed with log(x+1). Components retained for interpretation were
chosen according to the broken-stick model, in which only the components exceeding
the pattern of expected eigenvalues were considered significant (Jackson, 1993). Mouth
position and barbels were used as categorical variables. These analyzes were performed
in Rstudio using the BiodiversityR package (R Development Core Team, 2011).

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 Fish ecomorphology in oligotrophic streams

TABLE 5 | Contribution of the ecomorphological indices in the first three axes of the Principal
Components Analysis (PCA). Indices with higher correlations values for each principal component are
shown in bold and were selected for interpretation. See Tab. 3 for the list ecomorphological indices
abbreviations.

Ecomorphological index PC 1 PC 2 PC 3

Com_ind -0.799 -0.11 0.441

De_ind -0.934 -0.065 -0.127

Re_dep -0.884 -0.071 0.008

Rel_eye_si -0.913 0.053 -0.302

Eye_pos 0.948 0.088 -0.093

Re_area_pfin 0.287 -0.87 -0.214

As_ra_pfin -0.749 0.431 0.388

Ca_pen_tro -0.438 -0.345 -0.207

As_ra_cfin -0.559 -0.423 0.278

Re_area_cfin -0.248 -0.88 -0.057

Ca_pen_comin -0.59 -0.357 0.596

Re_len_capend 0.942 0.099 -0.083

Re_hig_Capend 0.736 0.267 -0.07

Re_wid_capend -0.537 -0.16 0.374

Re_area_dfin -0.07 -0.66 -0.123

Re_area_afin -0.849 0.044 -0.034

As_ra_afin -0.682 0.272 0.184

Re_area_pecfin 0.334 -0.819 -0.121

As_ra_pecfin -0.316 0.727 0.28

Re_hehead -0.601 -0.222 -0.571

Re_wihead -0.16 0.373 -0.774

Re_hemou -0.649 0.082 -0.592

Re_wimou -0.485 0.303 -0.755

O_gap_si -0.383 -0.026 -0.847

O_gap_sh -0.807 0.144 -0.029

Barb 0.955 0.026 -0.073

Mo_pos 0.814 0.021 -0.172

Eigenvalue 7.25 5.35 4.42

Explained variance (%) 44.9 16.6 11.4

Accumulated variance (%) 44.9 61.5 72.9

Broken-stick percentage 14.41 10.7 8.85

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Francisco J. Luque and Gabriel J. Colorado Z.

RESULTS

A total of 3,598 specimens belonging to 62 fish species, 27 families, and five orders
were captured (Tab. S1). Of the 34 non-rare species retained for the analysis, we
measured a total 472 individuals, where Characiformes was the dominant order
followed by Siluriformes, Cichliformes, and Cyprinodontiformes, respectively. With
respect to trophic guilds, nine species were classified as insectivores, eight omnivores,
six piscivores, six invertivores, and five detritivores (Tab. 3).
Mantel test showed that the morphological pattern of the fish assemblage had a
strong association with its phylogeny structure (r = 0.49; p = 0.001), habitat use (r =
0.18; p = 0.003) and trophic guilds (r = 0.296; p = 0.001), and a lack of association with
distance to river (r = -0.05; p = 0.835). After controlling for phylogenetic relationships,
partial Mantel tests showed that fish morphology was a good predictor of both trophic
structure (r = 0.227; p = 0.001) and habitat structure (r = 0.24; p = 0.025) in oligotrophic
headwater streams of northwestern Amazonia.
The first two components of PCA explained 60.9 % of total variation in the
ecomorphospace (Fig. 3). The ecomorphological traits with greater influence in
morphospace were related to feeding and locomotion (PC1, 44.9% of the variation)
and habitat use (PC2, 16%). PC1 shows a gradient, with positive loads corresponding
depressed bodies, dorsal eyes, small eyes, ventral mouth position, presence of barbels,
scores located towards the positive side. These characteristics categorized species as
benthic consumers of periphyton or invertebrates; this group was mostly composed
by the order of Siluriformes and families such as Trichomycteridae, Heptapteridae, and
Loricariidae. Whereas negative scores were associated to laterally compressed bodies,
larger anal fins, larger eyes, lateral eyes, and superior and subterminal mouths. These
characteristics are related to higher maneuverability and mobility, nektonic habits,
and tendency to chase prey such as fish and feed on insects and invertebrates adrift in
the water column. These traits are characteristics of characids, cichlids, and cyprinids
(Fig. 3). Likewise, PC2 shows a gradient as well. Positive scores were related to the
development of pectoral fins, characteristics of benthic species whereas negative scores
related to the development of dorsal and caudal fins, characteristics of active swimmers
or species that perform explosive movements.

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 Fish ecomorphology in oligotrophic streams

FIGURE 3 | Distribution of 34 fish species in the multivariate ecomorphological space generated by the first two PCA axes (above) and
the corresponding contribution of ecomorphological indices according to their scores (below). See Tabs. 2–3 for the list of species and
ecomorphological indices abbreviations, correspondingly.

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Francisco J. Luque and Gabriel J. Colorado Z.

DISCUSSION

Ecomorphology theory states that species morphological attributes are correlated with
their ecology and environmental factors, therefore it can act as predictor of how species
use several resources such as habitat (e.g., occupation in water column), and diet (e.g.,
foraging behavior) (Casatti, Castro, 2006; Soares et al., 2013). However, few studies
have evaluated the predictive ability of this theory in oligotrophic headwater streams
of northwestern Amazonia. In this study, habitat use and trophic structure attributes
showed a significant correlation with species morphology, while controlling for
phylogenetic effects. This suggests that fish assemblage in these oligotrophic headwater
streams is ecomorphologically structured, allowing to predict information about
habitat and trophic guild based on morphological features, e.g., if two species with
similar morphology were sampled, independently of their phylogeny, there is a great
probability that they would share same space (habitat) and food resources (trophic guild)
(Oliveira et al., 2010). Additionally, it indicates the high capacity of the used indices to
describe the trophic differences between morphologically similar species.
Studies evidencing the correlation between trophic ecology and morphology are
ambiguous. On one hand, Pouilly et al. (2003), Ward-Campbell et al. (2005), Ferreira
(2007;), Oliveira et al. (2010), found significant relationships between morphological
traits and diet, whereas Casatti, Castro (2006), Soares et al. (2013), Conde-Saldaña et al.
(2017) did not find any significance on these associations. This ambiguity might result
from the lack of capacity from several ecomorphological indices to reflect diet and
trophic differences among morphological similar species, trophic differences between
morphological similar species (Conde-Saldaña et al., 2017), and lack of standardized
indices protocols which resulted in researchers measuring same indices in different ways
(Villéger et al., 2017). Our results showed significant relationships between anatomy
and trophic structure and habitat use, indicating that fish assemblage is not defined by
phylogenetic relationships among species nor is randomly structured. However, we
highlight that due to our species selection criteria, our analyses only retained nearly half
of the species found in the sampled streams (34 out of 62) which, in turn, could have
influenced the obtained results. For instance, the input of morphological and ecological
different species captured but excluded from our analysis such as Potamorrhaphis guianensis
(Jardine, 1843), Eigenmannia virescens (Valenciennes, 1836), Otocinclus huaorani Schaefer,
1997, and Electrophorus electricus (Linnaeus, 1766) might help to better elucidate and
strengthen or reject the patterns found.
Our results failed to detect a relationship between the functional patterns (e.g.,
morphology and trophic guild) of the fish assemblage and distance to large, source
rivers. Several studies have found that species richness increase with distance from source,
which in turn increase trophic structure complexity, whereas headwater communities
showed simpler structures (Vannote et al., 1980; Bistoni, Hued, 2002; Fialho et al.,
2007) due mainly to the lack of habitat and resources diversity that increase as streams
increases their size. Our finding could result from the short distances these streams flow
until they reach the main river, Guaviare River, limiting the turnover of fish species
(Stegmann et al., 2019). The furthest sample point, Yam_int (Tab. 1), was only 30 km
from the Guaviare River with an altitudinal change of 81 m, and only displayed changes
in river substratum when compared to the other two sites downstream (FJL, pers. obs.).

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 Fish ecomorphology in oligotrophic streams

Most fish assemblages were reported to change after 150 km or when streams surpasses
fourth order (Grenouillet et al., 2004; Fialho et al., 2007). Further studies including
landscape position of samples point, habitat features, and other environmental variables
are needed to fully understand how these oligotrophic species rich and diverse streams
change along longitudinal gradient and within the dendritic network.
In this research, we identified two ecomorphological diversification patterns of
headwater streams of northwestern Amazonia based on interspecific differences in
ecomorphological traits. The first one is associated with feeding and habitat use. In
this context, two groups were observed in the ecomorphological space: 1) piscivorous,
omnivores, and insectivores, nektonic species with compressed bodies and 2)
detritivores and invertivores, benthic species with depressed bodies. Species of the first
group were associated with high maneuverability, larger lateral eyes, developed pectoral
fins, larger anal fins large mouths and heads, and compressed bodies. These species use
their vision to capture their prey and are good swimmers capable of stabilization of
movement (Watson, Balon, 1984; Breda et al., 2005; Conde-Saldaña et al., 2017). For
instance, omnivores and insectivores such as Anablepsoides sp., A. bimaculatus and B.
giacopinii have small dorsal fins, large eyes and pelvic fins which allowed them to move
throughout the water column, use different habitats and consume diverse food items
such as insects, fish, leaves, seeds, and fruits (Casatti, Castro, 2006; Conde-Saldaña et
al., 2017). Piscivores such as H. malabaricus and A. falcatus, had fusiform bodies that
allow high speed and larger heads and mouths, typical of ambush predators that wait
in marginal areas and dart to capture their prey (Breda et al., 2005; Ferreira, 2007). The
second group, detritivores and invertivores, have developed pectoral fins and caudal
peduncles, both fundamental for body stability on streams substrate and short distance
movements. Within this group, catfishes from families Loricariidae, Auchenipteridae,
and Trichomycteridae were most abundant and had depressed bodies, inferior sucker
mouths and dorsal eyes, typical of fishes that exploit intense current habitats (Ferreira,
2007; Oliveira et al., 2010; Prado et al., 2016). In turn, Cichliformes exhibit smaller
pectoral fins which provide better stability (Breda et al., 2005) for the use of structurally
complex habitats with slow moving waters such as macrophyte stands, roots and
backwaters (Breda et al., 2005; Oliveira et al., 2010; Conde-Saldaña et al., 2017).
The second ecomorphological diversification pattern was associated with locomotion.
This trend showed a gradient influenced by dorsal fin. Dorsal fin orientate motion as
it helps to avoid the rotation of fish along its longitudinal axis (Watson, Balon, 1984;
Breda et al., 2005). On one hand, loricariids use this fin as a keel and, together with their
developed pectoral fins, helps to keep the body on the bottom, whereas characids use
their small dorsal fin and elongated pectoral fins for continuous swimming throughout
the water column (Casatti, Castro, 2006). On the other hand, the large two dorsal fins
of cichlids greatly improve maneuverability (Breda et al., 2005).
Ecomorphology is a tool that can be used to explore intraspecific and interspecific
relationships and variations required to comprehend and predict complex relationships
between morphology and ecological specializations and generalizations (Soares et al.,
2013; Prado et al., 2016). Ecomorpholgy can be use to understand how fish assemblages
are affected by environmental changes such as logging, inflow change, and habitat
degradation (Lombarte et al., 2012; Jacob et al., 2021). This is particularly necessary
in the northwestern Amazonia as anthropogenic threats, including deforestation and

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Francisco J. Luque and Gabriel J. Colorado Z.

expansion of the agricultural frontier (Armenteras et al., 2013, 2019; Armenteras,

Rodríguez Eraso, 2014), cattle ranching (Etter et al., 2006a,b), illicit crops (Dávalos et
al., 2011), have increased since peace agreements ended armed conflict in Colombia
(Bogota-Gregory et al., 2024). These threats can pose strong deleterious effects on
fish fauna by altering streams integrity (Barletta et al., 2010; Pelicice et al., 2017). Our
results showed that fish assemblages in headwater oligotrophic streams of northwestern
Amazonia are spatially and trophically structured, meaning assemblages could be
negatively affected by changes in habitat structure along the streams. Thus, future
studies in these systems must incorporate the effect of anthropogenic disturbance on the
structure and functionality of fish assemblages.
This study provides novel information that can be used for effective management
and conservation efforts in this highly threatened region of the Amazonia, since it
contributes to understanding how fish assemblages are structured and how they can
be affected by current anthropogenic activities. Also, our results highlighted the use of
morphological traits as a tool to understand fish assemblage patterns and the ecological
interactions that affect them and their dynamics (Lombarte et al., 2012). Further studies
should include more species; analyze new possible ecomorphological patterns that
provide links between global changes, functional diversity and ecosystems services
(e.g., mediation of nutrient cycling through excretion and egestion, results that can be
associated to body mass using allometric coefficients). Additionally, the development
of a phylogenetic tree to better differentiate species phylogenetic relatedness
overcoming the weakness that might render the approach used in this research. Finally,
we recommend that ecomorphology should be incorporated into the research of fish
communities and advocate for a standardized protocols to avoid measuring same indices
in different ways, promoting consistency, and strengthen ecomorphology hypotheses.

ACKNOWLEDGMENTS

We thank Tania González, Juan David González, and Fabian Arias for their suggestions
and help with the graphical output, and Marco Melo and Edilberto Pachon for their
help during fieldwork. Funding was provided by the Cohort I Bicentennial Doctoral
Excellence Scholarship from The Ministry of Science, Technology, and Innovation of
Colombia (MINCIENCIAS) and the General Royalties System (Guaviare Department),
The Rufford foundation small grant (36352–1). We wish to thank IdeaWild for the
equipment donation. We express our gratitude to Universidad Nacional de Colombia.
Finally, we thank the reviewers for their comments and suggestions that greatly
improved our manuscript.

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 Fish ecomorphology in oligotrophic streams

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AUTHORS’ CONTRIBUTION
Francisco Javier Luque: Conceptualization, Data curation, Formal analysis, Funding acquisition,
Investigation, Methodology, Writing-original draft, Writing-review and editing.
Gabriel J. Colorado Z.: Conceptualization, Data curation, Formal analysis, Funding acquisition,
Investigation, Methodology, Writing-original draft, Writing-review and editing.

ETHICAL STATEMENT
This research was conducted under the scientific research permit No 0255, March 14, 2014, of the Autoridad
Nacional de Licencias Ambientales (ANLA) and approved by the Ethical Committee of the Facultad de
Ciencias of Universidad Nacional de Colombia, Bogotá.

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Diversity and Distributions Published by SBI HOW TO CITE THIS ARTICLE
• Luque FJ, Colorado Z. GJ. Spatial and trophic structure drive ecomorphological patterns
of fish assemblages in oligotrophic headwater streams of northwestern Amazonia. Neotrop
Ichthyol. 2025; 23(2):e230135. [Link]

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