Environ Biol Fish (2023) 106:1233–1246

https://doi.org/10.1007/s10641-023-01409-x

Morphological variation of African snakehead (Parachanna

obscura) populations along climate and habitat gradients
in Côte d’Ivoire, West Africa
Amien Isaac Amoutchi ·
Ogechi Nnabuchi Ugbor ·
Essetchi Paul Kouamelan · Thomas Mehner

Received: 12 October 2022 / Accepted: 18 March 2023 / Published online: 22 April 2023
© The Author(s) 2023

Abstract Intraspecific morphological variation may morphometrics of 15 fish body landmarks. Correla-
reflect phenotypic plasticity or adaptive divergence. tions between shape variations among populations
While adaptive shape divergence may occur more and climate and habitat characteristics and between
likely among isolated populations with reduced gene morphological and geographic distances were calcu-
flow, phenotypic plasticity may reflect morphologi- lated. We found significant morphological variations
cal responses to heterogeneous environments, even among the seven populations. The variations in fish
in spatially connected populations. We evaluated shape were concentrated on landmarks related to
both processes while examining morphological vari- swimming and feeding, suggesting a contribution of
ations among seven wild populations of snakehead environmental variation to morphological differentia-
fish (Parachanna obscura) along climate and habitat tion. However, we did not detect significant effects of
gradients in Côte d’Ivoire, West Africa. Morphologi- climate and habitat variables on fish shape. The trend
cal variations were studied by multivariate canoni- between geographical and morphological distances
cal variate analysis (CVA) as based on geometric was likewise not significant. Therefore, a mechanistic
understanding of the factors causing shape variation
among P. obscura populations in West Africa could
A. I. Amoutchi (*) · O. N. Ugbor not yet be achieved.
West African Science Service Centre On Climate Change
and Adapted Land Use (WASCAL) Graduate Research
Keywords Phenotypic plasticity · Environment ·
Program On Climate Change and Biodiversity, Université
Felix Houphouet-Boigny, Abidjan, Côte d’Ivoire Geographical distance · Geometric morphometrics
e-mail: amoutchiamienisaac@gmail.com

A. I. Amoutchi · T. Mehner
Introduction
Leibniz-Institute of Freshwater Ecology and Inland
Fisheries, Berlin, Germany
In Africa, many researchers have reported that
O. N. Ugbor freshwater organisms, especially fish populations, are
Department of Fisheries and Aquatic Resources
strongly affected by numerous threats including climate
Management, Michael Okpara University of Agriculture
Umudike, Umuahia, Nigeria change and anthropogenic perturbations such as dam
construction, water extraction, habitat modification,
E. P. Kouamelan water pollution, and overexploitation (Darwall et al.
Laboratoire d’Hydrobiologie, UFR Biosciences, Université
2011). For example, in Côte d’Ivoire, the climate has
Felix Houphouët Boigny, 22 BP 582 Abidjan 22, Abidjan,
Côte d’Ivoire significantly changed over the twentieth century,

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and the trend is expected to continue in the future. environment when future conditions can reliably be
Experts on The Intergovernmental Panel on Climate predicted (Sultan and Spencer 2002).
Change predicted an increase in temperature by 3 °C Cote d’Ivoire is a West African country charac-
by 2100, and a decrease of precipitation by 8% during terized by a diversified climate and water network
the next hundred years, under the RCP4.5 scenario in system. The climate divides the country along the
Côte d’Ivoire (Bernard 2014). It is predicted that by latitude into three principal climatic zones such as
the 2050s, hydrologic conditions that are significantly Guinean in the south, Sudano-Guinean in the mid-
different from the existing conditions will be dle, and Sudanian in the northern region. In addition,
experienced by more than 80% of Africa’s freshwater there is a fourth zone being the particular climate of
fish species (Darwall et al. 2011), which is likely to the mountain zone in the western region (Amoutchi
have a significant impact on fish diversity. et al. 2021). The Guinean zone is characterized by a
One of the African freshwater fish species with sub-equatorial climate, the Sudano-Guinean zone
great economic value is Parachanna obscura by an equatorial transition climate, and the Sudanian
(family Channidae), commonly known as African zone is characterized by a tropical climate. The water
snakehead fish. It is widespread in the intertropi- systems of the country are characterized by a vast and
cal convergence zone in Africa, where the water complex system including many lakes, wetlands, riv-
temperature ranges from 26 to 28 °C. Some stud- ers, and streams. Four major river basins with several
ies have indicated a decline in the populations of tributaries, namely, Comoe, Sassandra, Cavally, and
African snakehead fish, mainly due to overfishing Bandama, and many further coastal rivers are found in
and other anthropogenic threats coupled with the the country (Fig. 1) (Girard et al. 1970). Thus, if mor-
effect of the ongoing climate change (Lalèyè 2020; phological variation among P. obscura populations is
Amoutchi et al. 2021). However, Channidae spe- caused by responses to differing local environmental
cies, including P. obscura, can withstand stressful conditions and magnified by spatial isolation among
conditions, for example, low oxygen concentra- populations, we would expect a strong morphological
tions, notably by the possession of a cavity above variation of this species within Cote d’Ivoire.
the gill chamber, which functions as an accessory In our study, we used geometric morphometrics
respiratory organ (Lee and Ng 1991; Herborg et al. to examine morphological variation among seven
2007; Kpogue et al. 2012). In addition to these spe- wild populations of P. obscura along the climate and
cific organs acquired during phylogeny, variation in habitat gradients in Côte d’Ivoire. We tested whether
body morphology along climate and habitat gradi- shape variation correlates to differences of climate
ents (Munday et al. 2008; Lema et al. 2019; Del Rio and habitat characteristics and examined whether
et al. 2019) may increase the resilience of species to geographical distance among populations correlates
the effects of global change. with morphological distance. We expected the vari-
Intraspecific morphological variation is often pro- ations to be localized primarily in fish body parts
duced and maintained by divergent selective regimes, associated with ecological functions, for example, in
for example, in response to abiotic and biotic factors fins and caudal peduncle involved in swimming and
that differ among the habitats of populations. Under maneuvring, thus reflecting phenotypic plasticity. A
these conditions, morphological variation can reflect strong correlation between morphological and spatial
phenotypic plasticity of a common gene pool. How- distance might be indicative of a stronger effect of
ever, morphological diversity may also reflect adap- spatial isolation on morphological divergence.
tive processes resulting in strong genetic divergence
among populations (Lazzarotto et al. 2017), with the
potential to enable populations to reach new adap- Methods
tive peaks (Mouseau et al. 2000). While adaptive
morphological variation based on genetic divergence Field sampling and data collection
develops and is maintained more likely among spa-
tially isolated populations with reduced gene flow The study was conducted on several Ivorian inland
(Rundle and Nosil 2005), phenotypic plasticity is pre- watersheds (Fig. 1, Table 1). Bia River (KRIN), Sas-
dicted to be most advantageous in a heterogeneous sandra River (SBR), and San-Pedro Lake (SANP)

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Fig. 1  Map of the study area, showing locations and abbreviations of populations (Table 1) from which P. obscura specimens were
collected, and position of Côte d’Ivoire in West Africa

Table 1  Standard length (SL, cm), weight (g), and sample size from seven populations of P. obscura used in the study
River basin Watershed Population Sample Standard length (SL) Weight
Size (N) Range (cm) Mean ± SE Mean ± SE

Niger River Bague River BAG 22 14.5–37.8 24.5 ± 6.2 208 ± 164
Bandama Kan Lake KAN 18 19.4–37.6 27.7 ± 5.5 269 ± 155
Coastal River Bia Bia River KRIN 12 16.2–35.1 20.9 ± 4.7 131 ± 136
Sassandra Nzo River NZO 40 16.5–28.5 21.4 ± 3.4 138 ± 74
Coastal River San-Pedro San-Pedro Lake SANP 26 18.4–32.8 25.5 ± 4.9 254 ± 139
Sassandra Buyo Lake SAS 32 14.7–25.5 20.3 ± 2.9 108 ± 39
Sassandra Sassandra River SBR 17 22.9–31.0 27.6 ± 2.5 297 ± 81

were sampled in the southern region (Guinean cli- Bagoue River (BAG) from the northern region was
mate zone). Kan Lake (KAN), located centrally, and sampled in the Sudanean climate zone.
Buyo Lake (SAS) and Nzo River (NZO), located in A total of 167 fish from seven P. obscura popu-
the center-western part of Cote d’Ivoire, were sam- lations were collected (Table 1). P. obscura fisher-
pled from the Sudano-Guinean zone. Finally, the men from each sampling site were employed for fish

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sample collection using traps and fish nets. Within Morphological data collection
1 h from capture, the left side of each specimen was
photographed using Nikon (D5600) camera (cali- We digitized 15 landmarks (LMs) on 2D pictures to
brated against a graduated ruler to standardize meas- obtain the x–y coordinates of all points using Tps-
urements) for geometric morphometric analysis. The Dig2 v31 (Rohlf 2015) (Fig. 2).
short phase between capturing and photographing
fish avoided unwanted morphological distortions, for Data analysis
example, from rigor mortis. Specimens’ length and
weight were also measured. Then, specimens were Morphological variation
fixed in 10% formaldehyde solution and transported
to the laboratory for further analysis. To perform geometric morphometric analysis, land-
Physico-chemical parameters of each site includ- marks were converted to shape coordinates by the
ing water temperature (°C), pH, total dissolved generalized least square Procrustes superimposition
solids (ppm), electrical conductivity (µS/cm), and (GPA; Rohlf and Slice 1990). This method transforms
redox potential (mV) were measured in situ with landmark configuration by preserving all informa-
a handheld multi-parameter monitoring device tion about shape differences among specimens but
(WQM-243/PHTK-243, TEKCOPLUS, Hong Kong, removes non-shape variability in raw coordinates
China). Water depth (cm) and water transparency such as location, orientation, scale, and rotation, and
(cm) were also measured (Appendix Table 4). The then standardizes each specimen to unit centroid size
canopy cover of vegetation (%) along the shore- (CS). Centroid size represents a dimensionless size
line was estimated visually. Measurements were measure computed as the square root of the summed
done over the dry and rainy seasons at each site in squared Euclidean distances from each landmark to
September 2020 and January, June, and December the specimen centroid (Bravi et al. 2013). The shape
2021. The values were later averaged to character- data obtained were further size-corrected using a
ize the situation of the locations during both seasons regression of shape (Procrustes coordinates) on size
(Appendix Table 4 ). The latitude and longitude of (log centroid size) for each population separately. The
each site were measured using a handheld global obtained regression residuals were used as standard-
positioning system. Additional climate data (mean ized shape data in further analyses on shape varia-
air temperature, maximum and minimum air tem- tions (see for a similar approach Scharnweber et al.
perature, relative humidity, and mean precipitation) 2013). By using a set of 21 polymorphic microsatel-
were obtained for each sampling site from NASA lite markers, we have recently demonstrated that the
POWER Data Access (https://​power.​larc.​nasa.​gov/​ populations of snakehead fish in Côte d’Ivoire includ-
data-​access-​viewer/) by entering the latitude and ing those studied here were genetically differenti-
longitude of sampling sites (Appendix Table 4 ). ated (Amoutchi et al. 2023). Therefore, we applied

Fig. 2  Defined landmark points for collection of P. obscura caudal fin, 8—posterior end of the medial region of the caudal
body shape data. Landmarks are1—anterior-most point of the peduncle, 9—posterio-ventral end of the caudal peduncle at
snout tip on the upper jaw, 2—anterior-point of nasal bones, its connection to the caudal fin, 10—insertion and 11—origin
3—center of the eye, 4—superior margin of the head, 5—ori- point of the anal-fin base, 12—most anterior point of pelvic
gin and 6—insertion point of the dorsal-fin base, 7—poste- fin, 13—most anterior point of pectoral fin, 14—posterior edge
rio-dorsal end of the caudal peduncle at its connection to the of the opercule, and 15—ventral end of the gill slit

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canonical variate analysis (CVA), a multivariate sta- Results

tistical method used to distinguish variation among
multiple pre-defined groups (populations) of speci- Morphological variation among P. obscura
mens, to quantify inter-population shape variations. populations
CVA results are statistically reported as Mahalanobis
distance (Md), which represents a measure of dis- Average length and weight of specimens varied
tance among populations relative to the within-pop- among populations (Table 1). Fishes of KAN and
ulation variation. The statistical significance among SBR were the largest, while NZO, KRIN, and SAS
Mahalanobis distances were determined by bootstrap- populations were composed of smaller specimens.
ping the Md matrix (permutation test with 10,000 However, the length range of specimens suggested
runs). All the subsequent analyses were performed in that only adult individuals were included in the study.
MorphoJ version 1.07a software. Probabilities < 0.05 Canonical variate analysis indicated significant
were considered significant. Group assignments were (p < 0.05) Mahalanobis distances (Md) among all popu-
cross-validated by a jackknife resampling test, per- lation pairs (Appendix Table 5 ), with the highest Md
formed in R package Morpho (Schlager 2017). values observed between KRIN, KAN, and the other
populations. The classification rates obtained from
the CVA based on the jackknife cross-validation test
Influence of geographical isolation or distance
resulted in an overall classification rate of 60% of the
on morphological variations
specimens into their original groups. The highest classi-
fication rates were recorded for NZO and SANP, while
To test for the effect of geographic distance on the
the lowest classification rate was recorded for BAG
morphological variation among P. obscura popula-
(Table 2), indicating high intra-group variability within
tions, the Mantel correlation between the pairwise
this population. The first two CVA factors contributed
Mahalanobis distances (Appendix Table 5) and the
38.4% (CV1) and 21.9% (CV2) of the total variation.
natural logarithm (ln) of the pairwise Euclidean geo-
CV1 reflected variation in the caudal body part (land-
graphic distances (km) was calculated. The geograph-
marks 6, 7, 8, 9, and 10), dorsal fin length (landmark
ical distance matrix was calculated by QGIS 3.22.6
5), and upper body depth (Fig. 3). CV2 described vari-
using the global positioning system (GPS) coordi-
ation in the anterior region of the body, notably in the
nates of the sampling locations. Mantel correlation
snout orientation (landmarks 1 and 2), lower body
was run in vegan package (Oksanen et al. 2022) in R,
depth (landmark 11), the shape of the upper region
with 1000 permutations.
of the head (landmark 4), eyes position (landmark 3),
and the positions of the pectoral fin and posterior edge
Climate and habitat variability and its influence on P. of opercula (landmarks 13, 14). Overall, however, the
obscura morphology morphological variation between individuals and popu-
lations was relatively weak, and in no region of the fish
To assess the environmental heterogeneity among body was particularly strongly accentuated.
collection sites of P. obscura populations, principal Despite the statistically significant differentiation
component analysis (PCA) on all physico-chemical, among populations, individuals of KAN and BAG
climatic, and habitat variables (Appendix Table 5 ) were strongly scattered in the CVA plot, and hence,
was applied. The PCA was calculated by R.4.1.2 soft- their population ellipses overlapped substantially
ware using the factoextra’ package (Kassambara and with those of other populations (Fig. 4). Apart from
Mundt 2020). For testing the link between fish body these two populations, the other five populations were
shape and climate and habitat variables, Pearson cor- grouped distinctly from each other on the scatterplot.
relations were calculated between the first two prin- NZO and SANP were the most distinct groups, with
cipal components explaining the highest percentage only few individuals overlapping with other groups.
of variability in climate and habitat variables among Individuals of KAN and KRIN populations with nega-
sampling sites and the population centroids of body tive CV1 and CV2 values had a deeper and slightly
shape from CVA. Pearson correlation was computed shorter caudal peduncle and dorsal fin, less deep body,
in the stats package in R4.1.2 (R Core Team 2021). a posterior edge of their operculum and pectoral fin in a

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Table 2  Number of specimens classified into the seven popu- of canonical variate analysis. N represents the total number of
lations of snakehead fish, and percentage of correctly classified specimens in each population
specimens, as obtained from a jackknife cross-validation test
N BAG KAN KRIN NZO SANP SAS SBR

BAG 22 8 (36%) 1 2 3 3 4 1
KAN 18 3 11 (61%) 1 0 1 1 1
KRIN 12 1 2 6 (50%) 1 0 1 1
NZO 40 1 0 1 28 (70%) 4 3 3
SANP 26 2 0 0 4 18 (69%) 1 1
SAS 32 4 0 2 4 1 18 (56%) 3
SBR 17 2 0 1 2 1 1 10 (59%)

Fig. 3  P. obscura shape

variations corresponding
to wireframe deformation
along the first two CVA
axes (CV1, CV2). Grey
shape lines are the mean
shapes of all specimens, +2
and black lines are shape CV1
modifications along the two
-2
main CVA axes. The black
dots numbered in red reflect
the 15 landmarks measured

+2
CV2
-2

downwardly sloping position, the snout slightly sloping posterior edge of their operculum and pectoral fin in a
upwards, and eyes positioned in the bottom part of the downwardly sloping position, the snout slightly sloping
head towards the snout, in contrast to individuals from upwards, and eyes positioned in the bottom part of the
NZO. Individuals of SANP with negative CV2 and head towards the snout. These individuals were mor-
positive CV1 values were characterized by a slightly phologically opposed to individuals of the SAS popu-
longer caudal peduncle narrowing at the insertion bases lation. Finally, SBR individuals, with negative CV2
of the dorsal and anal fins, a slightly longer dorsal fin, a values and central CV1 values, had a posterior edge of

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Fig. 4  Scatter plot obtained of first two factors of the canoni- as an ellipse (containing 80% of the individuals in the popula-
cal variate analysis on 15 geometric morphometrics characters tion in question), and their centroid (mean) as a large central
(landmarks). Individuals of different populations are repre- symbol
sented by a different set of symbols. Populations are presented

their operculum and pectoral fin in a downwardly slop- total variation (Fig. 6). PC1 (47.8% of total varia-
ing position and the snout slightly sloping upwards with tion) mainly reflected the variation in climatic vari-
eyes positioned in the bottom part of the head towards ables such as relative humidity (RH), maximum
the snout. Individuals of the BAG population were air temperature (T_max), mean air temperature
morphologically strongly diverse, and hence, their posi- (T_Mean), minimum air temperature (T_min) and
tions were scattered throughout the diagram. mean precipitation (precipit), as well as the habi-
tat variables like water temperature, redox poten-
Influence of geographical distance on morphological tial (ORP), electrical conductivity (cond), and total
variations dissolved solids (TDS). T_max, T_Mean, ORP,
and water temperature were negatively correlated
There was no significant correlation between pair- with PC1, while all other variables were positively
wise morphological distance and geographical dis- correlated with PC1 (Table 3). PC2, with a load-
tance among the seven populations (Mantel test, ing value of 28.5%, mainly reflected the variation
P = 0.23, Fig. 5). of habitat variables, including water depth (Depth),
transparency (Transp), and canopy. Except canopy,
which was positively correlated with PC2, all other
Climatic and habitat characteristics of collection sites variables were negatively correlated with PC2. The
of P. obscura populations habitat of the KRIN population was characterized
by a high canopy, low water transparency, acidic
PCA of 13 climate and habitat variables extracted pH, and shallow water depth. The habitats of NZO,
two principal components explaining 76.2% of the SAS, and SBR populations were characterized by

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Fig. 5  Estimation of
pairwise morphological
distances between popula-
tions plotted against natural
logarithm (ln) of pairwise
Euclidean geographical dis-
tance (km). P was obtained
based on the Mantel test

Fig. 6  Scatterplot of the projection of the populations (popula- mum air temperature (°C); RH, relative humidity (%); precipit,
tions abbreviation see Table 1), and habitat and climate vari- mean precipitation (mm/day); T_water, water temperature
ables (arrows) on the first two principal components explain- (°C); Depth, water depth (cm); Cond, electrical conductivity
ing 76.2% of the total variation. T_Mean, mean air temperature (µS/cm); TDS, total dissolved solids (ppm); ORP, redox poten-
(°C); T_max, maximum air temperature (°C); T_min, mini- tial (mV); Transp, transparency (cm); Canopy, canopy (%)

high water transparency, neutral or basic pH, low populations had higher mean and maximum air
water and air temperature, a short canopy, moder- temperatures, higher water temperatures, high
ately high precipitation and humidity, and deeper redox potential, and low precipitation and relative
water. The collection sites for BAG and KAN humidity. The SANP population collection site was
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Table 3  Loadings of the 13 climate and habitat variables on Discussion

the first two principal components explaining 76.2% of total
variation and with Eigenvalues greater than 1. Highest loading
values are bolded
We analyzed intra-specific morphological variation
of P. obscura populations along climate and habitat
Variables PC1 PC2 gradients in freshwater habitats of Côte d’Ivoire. Sig-
T_Mean − 0.7 0.6 nificant morphological variation was observed among
T_max − 0.9 0.0 all populations, although the within-population vari-
T_min 0.8 0.4 ability was also high, and was particularly strongly
RH 0.9 − 0.1 expressed in the population with the lowest number of
Precipit 0.98 0.0 studied individuals (KRIN). The morphological vari-
T_water − 0.8 0.1 ations were principally documented in the head, cau-
Depth 0.4 − 0.9 dal peduncle, and pectoral and pelvic fins. Other stud-
pH 0.3 − 0.3 ies on fish species such as Gobiocypris rarus (Shao
Cond 0.7 0.6 et al. 2007), Hemigrammus coeruleus (Lazzarotto
TDS 0.7 0.6 et al. 2017), Nothobranchius orthonotus ( Bravi et al.
ORP − 0.9 − 0.2 2013; Vrtílek and Reichard 2016), Channa punctatus
Transp 0.1 − 0.7 (Kashyap et al. 2016), or other Cyprinidae family fish
Canopy − 0.2 0.9 (Jacquemin and Pyron 2016) corroborate our findings
that intra-specific morphological variation in fishes
T_Mean mean air temperature (°C), T_max maximum air tem-
perature (°C), T_min minimum air temperature (°C), RH rela-
can be strongly significant. However, we did not find
tive humidity (%), precipit mean precipitation (mm/day), T_ potential causes of intra-specific variability. The iso-
water water temperature (°C), Depth water depth (cm), Cond lation by distance test showed no significant correla-
electrical conductivity (µS/cm), TDS total dissolved solids tion between morphological variations and geograph-
(ppm), ORP redox potential (mV), Transp transparency (cm),
Canopy canopy coverage of what vegetation (%)
ical distance. Furthermore, there was only a weak
correspondence between morphological variation and
differences of environmental conditions among popu-
the most polluted one with the highest values for lations. Accordingly, we did not find significant evi-
chemical parameters such as electrical conductivity dence for environmental-driven phenotypic plasticity
and total dissolved solids. Precipitation and humid- or for the effects of spatial isolation on the expres-
ity were also higher at this site. sion of intraspecific morphological variation in this
species.
Correlation between morphological variation and CVA results showed that most shape variations
climate and habitat variables were localized in the head (shape, eye, and snout posi-
tions), caudal peduncle, and pectoral and pelvic fins.
Overall, there was only a weak correspondence The study by Mouludi et al. (2019) on two popula-
between morphological variation and environmental tions of Channa gachua, another species belonging to
conditions among populations (Fig. 7). The high- the Channidae family, also showed body shape differ-
est correlation coefficient (r = 0.68, P = 0.09, n = 7, ences in the position of the snout and caudal pedun-
df = 5) was obtained between CV1 and PC1. With cle length. Similar morphological variations mostly
declining temperature, increasing precipitation, localized in fins and head region were observed
humidity, and conductivity, and at higher productivity among populations of Lavinia symmetricus (Brown
(TDS) (high PC1), the caudal peduncle of snakehead et al. 1992). Variation in eye diameter and head length
fish was elongated and narrows at the insertion bases was observed among the population of Trachurus tra-
of the dorsal and anal fins, the dorsal fin becomes churus through linear morphometrics (Bektas and
longer, the body becomes deeper in its anterior part, Belduz 2009). A study on Phycis phycis populations
and the posterior edge of the operculum and pectoral also showed that morphological variation was mainly
fin shift towards a downwardly sloping position (high found in fin regions as well as in the position of the
CV1). All other combinations of PC and CV axes posterior limit of the operculum (Vieira et al. 2016).
were less strongly correlated. These regions of fish body are mainly associated with

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Fig. 7  Pearson correlation plots between the first two CVA bles between sites. Upper left plot—CV1 and PC1, upper right
axes (CV1 and CV2), representing morphological variation plot—CV1 and PC2, lower left plot—CV2 and PC1, and lower
among populations, and the first two PCA axes (PC1 and right plot—CV2 and PC2. The correlation coefficient (r) and
PC2), which represent variation in habitat and climate varia- probability (p) are shown on each plot

functions such as fish locomotion (pelvic and pecto- of lower discharge. Scharnweber et al. (2013) dem-
ral fins and caudal peduncle) and feeding (eyes and onstrated that Rutilus rutilus cyprinids from lakes
snout), vital functions that are essential for the sur- with high predation pressure were characterized by a
vival of the species. Fin and caudal peduncle dimen- more streamlined body and caudally inserted dorsal
sions are related to swimming ability, which includes fin, attributes that facilitate escape from predators.
speed and maneuverability (Sampaio et al. 2013; Li Phenotypic plasticity of body morphology is regarded
et al. 2021). Thus, variation in these regions may as a key strategy for populations living in changing
help the fish coping with environmental variability environments, although morphological variation may
between different locations. even play a role in adaptive diversification (Svanbäck
However, in our study, no significant associations 2004). We did not find significant evidence of mor-
were found between fish shape and climate and habi- phological variations in the African snakehead fish
tat variables. Shape variation of many species can in response to variation in environmental conditions.
respond to temperature, dissolved oxygen, radiation, However, there was a relatively high correlation coef-
water depth, food availability, current or hydrol- ficient obtained between CV1 and PC1 (r = 0.68),
ogy, or correlated with their feeding mode, predation which supports that morphological variation between
risk, and habitat use (Swain et al. 2005; Bunje et al. snakehead fish populations may in part be affected by
2007; Spoljaric and Reimchen 2011; Antonucci et al. environmental variation. Thus, further investigations
2012; Bravi et al. 2013; Scharnweber et al. 2013; including more populations may shed light on the fac-
Jacquemin and Pyron 2016). Jacquemin and Pyron tors that induce intra-specific morphological variation
(2016) demonstrated that changes in the morphology in this fish species.
of Cyprinid fishes can be related to hydrology, where Geographical isolation of populations can lead to
more fusiform individuals with downward turned and morphometric variations between populations, and
extended caudal peduncle dominated during periods this morphometric variation can provide a basis for

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population differentiation (Bookstein 1991). Spatial fish shape and climate and habitat variables were
isolation by unconnected river systems or by con- weak, and spatial isolation likewise did not contribute
nectivity barriers within river systems may prevent substantially to the differences of fish shape among
gene flow among populations, and hence, adaptive populations. The differences in shape as obtained by
morphological diversification may have contrib- geometric morphometrics may reflect phenotypic var-
uted to shaping the morphological variation among iation that may aid the fish in surviving in a variety
populations. In contrast, connectivity of river sys- of environments and may also contribute to enhanced
tems facilitates the movement of organisms, nutri- fitness under the various threats of freshwater ecosys-
ent transfer, and energy flow across the landscape tems in Africa, such as climate change, gold mining,
(Opperman et al. 2010), and is used by fish for repro- water withdrawal for human needs, overexploitation,
ductive migrations, recruitment and recolonization, industrial waste discharge, pesticide use for agri-
and access to floodplains (Pettit et al. 2017). Spatial cultural purposes along watersheds, and obnoxious
connectivity also allows genetic connectivity among fishing practices (Amoutchi et al. 2021). However,
populations and would facilitate weak genetic diver- a functional and mechanistic understanding of the
gence, such that morphological variation would adaptive value of shape variations between popula-
reflect primarily phenotypic plasticity. In our study, tions would require common garden experiments and
no significant association was observed between extended analyses of functional genomics, data that
spatial and morphological distances, despite the fact are currently not yet available.
that we found significant morphological variations
(as indicated by Mahalanobis distances) among the Acknowledgements We would like to thank two anonymous
referees for their valuable comments that helped improving
populations obtained from hydraulically unconnected the text. We are grateful to WASCAL (sponsored by the Ger-
sites. Furthermore, we also found differentiation man Federal Ministry for Education and Research) and Leibniz
among populations from the same basin (SBR and Institute of Freshwater Ecology and Inland Fisheries (IGB) for
SAS, both from Sassandra River), likely caused by sponsoring the work.
a dam constructed on Sassandra River. Indeed, the
Author contribution Conceived and designed the investiga-
SBR population was collected from the downside of tion: AAI, TM, and EKP. Performed field and laboratory work:
the Sassandra River downstream of the dam, while AAI and ONU. Analyzed the data: AAI and ONU. Wrote the
the SAS population was collected from the upstream first draft of the manuscript: AAI. Guided the data analysis and
side of the river in the artificial Lake Buyo which is the structure of the manuscript: TM. Supervised the project,
reviewed and approved the final manuscript: TM and EKP. All
the reservoir impounded by the dam. Finally, fishes co-authors provided manuscript edits.
of the NZO population that were collected from Nzo
River, a tributary of Lake Buyo, were also morpho- Funding Open Access funding enabled and organized by
logically different to SBR and SAS populations. A Projekt DEAL.
significant effect of spatial isolation on genetic dif-
ferentiation of snakehead fish populations including Data availability The data generated during the current
study are available from the corresponding author upon reason-
those studied here (Amoutchi et al. 2023) supports able request.
that spatial arrangement of rivers and potential con-
nectivity among fish populations can be important. Declarations
Larger sample sizes and a spatially refined sampling
scheme be needed to detect the spatial effects on the Ethical approval All methodologies for conducting the activ-
morphometry of the snakehead fish. ities of this study were approved by the Ethics Committee of
WASCAL—Graduate Programme in Climate Change and Bio-
diversity, Université Felix Houphouet-Boigny, Côte d’Ivoire.
Authorization for data collection was also obtained from the
Conclusion Ivorian Ministry of Animal Resources and Fisheries.

The current study demonstrates significant morpho- Consent for publication All authors consent for the publica-
logical variation among seven P. obscura popula- tion of this work.
tions from sites that strongly differed in climate and Conflict of interest The authors declare no competing interests.
habitat variables. However, the correlations between

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Appendix Table 4 Table 5

Table 4  Climate and habitat variables from the locations of the seven P. obscura populations sampled. Values are averages from
measurements during the dry and wet seasons. Data origin is either from NASA database or by direct measurements (DM)
Population T_Mean T_max T_min RH precipit T_water Depth pH Cond TDS ORP Transp Canopy

BAG 26.2 40.2 11.4 65 3.4 28.4 171 6.9 41.7 27.2 281.4 42.5 50
KAN 26.7 39.4 14.8 75 3.5 30.8 222 7.2 118 68.5 271.2 65.0 48
KRIN 26.2 35.0 16.4 85 5 28.6 289 6.1 41.7 25 226.9 49.9 62
NZO 24.9 32.7 14.4 85 5.1 27.4 1273 7.1 67.5 45 276.3 36.8 37
SAS 25.2 36.2 14.4 82 4.8 28.9 1650 8.1 115 85 195.0 90.0 18
SBR 25.1 32.5 16.0 88 6 25.1 1607 7.0 96 67 197.2 98.5 23
SANP 25.6 31.7 18.3 87 6.5 26.2 142 7.6 1042 720.8 117.5 37.5 57
Origin NASA NASA NASA NASA NASA DM DM DM DM DM DM DM DM

T_Mean mean air temperature (°C), T_max maximum air temperature (°C), T_min minimum air temperature (°C), RH relative
humidity (%), precipit mean precipitation (mm/day), T_water water temperature (°C), Depth water depth (cm), Cond electrical con-
ductivity (µS/cm), TDS total dissolved solids (ppm), ORP redox potential (mV), Transp transparency (cm), Canopy canopy coverage
of what vegetation (%)

Table 5  Mahalanobis BAG KAN KRIN NZO SANP SAS SBR

distances (lower diagonal)
and significance (upper BAG < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001
diagonal) of pairwise
KAN 3.3 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001
comparisons by the
canonical variate analysis KRIN 3.2 4.1 < 0.0001 < 0.0001 < 0.0001 < 0.0001
performed between NZO 2.9 4.2 4.8 < 0.0001 < 0.0001 < 0.0001
populations of the P. SANP 3.5 4.5 5.1 3.3 < 0.0001 < 0.0001
obscura. Results were
SAS 2.3 3.4 3.5 3.1 4.0 < 0.0001
obtained from 10,000
permutation runs SBR 3.1 4.0 4.2 3.4 3.1 3.2

Open Access This article is licensed under a Creative Commons Amoutchi AI, Kersten P, Vogt A, Kohlmann K, Kouamelan EP,
Attribution 4.0 International License, which permits use, sharing, Mehner T (2023) Population genetics of the African snakehead
adaptation, distribution and reproduction in any medium or format, fish Parachanna obscura along West Africa’s water networks:
as long as you give appropriate credit to the original author(s) and implications for sustainable management and conservation.
the source, provide a link to the Creative Commons licence, and Ecol Evol 13:e9724. https://​doi.​org/​10.​1002/​ece3.​9724
indicate if changes were made. The images or other third party Antonucci F, Boglione C, Cerasari V, Caccia E, Costa C (2012)
material in this article are included in the article’s Creative Com- External shape analyses in Atherina boyeri (Risso, 1810)
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mons licence and your intended use is not permitted by statutory atlantic horse mackerel, Trachurus trachurus populations
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