International

Satour, Journal of...Odonatology

Hezil, Taferghoust Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria
2024, Vol. 27, pp. 172–186
doi:10.48156/1388.2024.1917289

Land use and beyond:

unraveling environmental determinants
of odonate assemblages in northeastern Algeria

Abdellatif Satour a,b, Wissem Hezil a,b

, Meriem Taferghoust a,b,
Hayat Boucenna a,b, Farrah Samraoui a,b
& Boudjéma Samraoui a,c,*

a
Laboratoire de Conservation des Zones humides. Université 8 Mai 1945 Guelma, Algeria
b
Department of Ecology. University 8 Mai 1945 Guelma, Algeria
c
Department of Biology. University Badji Mokhtar, Annaba, Algeria
*Corresponding author: Email: [email protected]

Abstract. Freshwater ecosystems, recognized as hotspots of biodiversity, are under in-

creasing threat from human activities. Odonata, the iconic inhabitants of these ecosys-
tems, are highly sensitive to changes in habitat morphology, hydrology and land-use dy-
namics. In this study, the odonate assemblages in four different regions in northeastern Al-
geria are investigated to identify the key environmental factors that influence them. These
regions differ in terms of climate, altitude and land use. Over the course of the study, we
documented twenty-one species, including ten Zygoptera and eleven Anisoptera, which
together account for a substantial proportion (~47%) of regional odonate diversity. Spe-
cies composition is correlated with habitat typology and land cover. Multivariate analyses
revealed distinct ecological patterns, highlighting streambed width, flow velocity and, to
a lesser extent, land cover and altitude as influential factors in assemblage segregation.
Research Article This study underscores the importance of flow velocity, which is known to facilitate oxygen
uptake by larvae and influence the distribution and composition of species. Altitude, a
OPEN ACCESS
key factor influencing phenology, growth and development, and land use (including for-
This article is distributed
under the terms of the
ests, urbanization, agriculture and pasture) are also identified as potential determinants
Creative Commons of odonate assemblages. To effectively tailor conservation strategies to specific regional
Attribution License, conditions around the world, the study emphasises the importance of identifying the key
which permits unrestricted use, environmental factors affecting odonate assemblages.
distribution, and reproduction in Key words. Odonata, freshwater ecosystems, hotspots, landscape cover, land use, North
any medium, provided the original Africa
author and source are credited.

Published: 30 July 2024

Received: 27 August 2023 Introduction
Accepted: 25 June 2024
The Mediterranean region, considered the cradle of biodiversity (Médail & Qué-
Citation: zel, 1997; Mittermeier et al., 2004; Myers et al., 2000), has provided indispens-
Satour, Hezil, Taferghoust, Boucen-
able ecosystem services to local communities for millennia (Blondel & Aronson,
na, Samraoui & Samraoui (2024):
Land use and beyond: Unraveling 1999). This region functions as a refuge for myriad endemic and threatened spe-
environmental determinants of cies (García et al., 2010; Riservato et al., 2009) that have adapted to the spa-
odonate assemblages in tial and temporal heterogeneity of the landscape, which is characterized by sea-
northeastern Algeria. sonal and predictable droughts and floods (Bonada et al., 2007; Gasith & Resh,
International Journal of 1999). Nevertheless, the wetlands in this region have felt the effects of increasing
Odonatology, 27, anthropogenic pressure, which has led to a sharp decline in the abundance of
172–186
macro­invertebrates and the areal extent of their habitats in recent years. (Bensli-
doi:10.48156/1388.2024.1917289
mane et al., 2019; Díaz-Paniagua & Aragonés, 2015; Taylor et al., 2021; Zalidis et
Data Availability Statement: al., 1997).
All relevant data are Among the Mediterranean countries, Algeria, located in North Africa, has a high
within the paper. number of wetlands of global importance. Many of these wetlands are designated

International Journal of Odonatology │ Volume 27 │ pp. 172–186 172

Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

as Ramsar sites, highlighting their central role in inter- velocity, substrate, helophytes, etc.) and the composi-
national conservation efforts (Samraoui & Samraoui, tion and diversity of odonates, which include a group
2008). Yet, amidst their ecological significance, their of threatened species. In particular, by assessing the
role as refuges for endangered fish, amphibians and effects of landscape cover and land use on odonate as-
aquatic insects has often been eclipsed by ornithologi- semblages, the study can help identify potential man-
cal considerations. While the importance of odonates agement strategies that can help protect and conserve
in Algerian wetlands, particularly in the El Kala region, these important organisms and their habitats in North
has been recognized since the 19th century (Sélys-Long- Africa.
champs, 1849, 1865), certain areas remain unexplored
despite published efforts (Samraoui & Corbet, 2000;
Samraoui & Menaï, 1999). Materials and methods
Dragonflies (Insecta: Odonata), often referred to as Study area description
“guardians of the watershed” (Clausnitzer & Jödicke,
2004), have special ecological significance, serving as The survey was conducted in northeastern Algeria (Fig-
indicators of the health of aquatic habitat and the ad- ure 1). It focused on four different regions, each char-
jacent environment (Chovanec, 2000; Ferreras-Romero acterized by different climatic regimes, from the humid
et al., 2009). However, it is important to note some conditions of Collo (Skikda) to the semi-arid environ-
odo­nates act as indicators of polluted areas (Bouhala et ments of Oum El Bouaghi, Batna, and Khenchela (Bou-
al., 2019; Campos et al., 2014). They also provide vari- cenna et al., 2023).
ous ecosystem services that have provisioning, cultural, Collo, an important town, is located 71 kilometers
supporting and regulatory functions and benefit hu- northeast of Skikda. It is nestled in the Atlas-Tellian
man society (Simaika & Samways, 2008). The intricate Mountains, which stretch from northwestern Tunisia
life cycles of Odonata provide critical linkages between to northeastern Morocco. The region has a Mediterra-
aquatic and terrestrial communities (Remsburg & Turn- nean climate with distinct seasons: a hot and dry sum-
er, 2009). mer and a cold and humid winter. The average annual
Due to anthropogenic changes, habitat loss and rainfall varies between 800 and 1400 mm, while the
degradation are widespread across the Earth’s surface average annual temperature is 20.2°C (Boudemagh et
(Ellis & Ramakatty, 2008; Smith, 2007). Despite their al., 2013). Sampling in the region was carried out at five
ecological value, freshwater ecosystems in which odo- river basins (Figure 2a).
nates thrive face major challenges worldwide (Abell, Oum el Bouaghi, a wetland complex, spans the east-
2002; Brautigam, 1999). These ecosystems are among ern Hauts Plateaux. This region forms a bridge between
the most endangered on Earth and are under severe the Mediterranean Sea and the Sahara Desert and in-
threat from habitat destruction, modification and pol- cludes mountains, valleys and plateaus. The climate is
lution due to factors such as drainage, dams, water semi-arid and is characterized by an average annual
withdrawals, pollution and invasive species (Dudgeon temperature of 17.51°C and an annual rainfall of less
et al., 2006; Malmqvist & Rundle, 2002). The resulting than 350 mm. The region is dominated by steppe veg-
impacts on biota have led to a rapid decline in the struc- etation and is interspersed with temporary salt lakes
tural integrity of aquatic and terrestrial ecosystems called “chotts”, “garaas” or “sebkhas”. Nine different
(Hallmann et al., 2017; Romero et al., 2021; Sánchez- wetlands were studied in this area, including chotts,
Bayo & Wyckhuys, 2019; Wagner, 2020). garaas, sebkhas, dams and four intermittent streams
A targeted research program aimed to understand (wadis or oueds) (Figure 2d).
the crucial role of odonates in North African wetlands The Aures Massif lies to the east amidst the foot-
and to develop targeted conservation strategies. This hills of the Sahara Atlas. The region lies between the
involved exploring the complex relationships between northeastern Hauts Plateaux and the southern Saharan
odonate assemblages and their environmental condi- Depression and has a wide climatic range. The north-
tions (Samraoui & Corbet, 2000). While the research ern slopes are characterized by cool winters with tem-
covered a significant part of northern Algeria, the fo- peratures below 4°C, while the southern slopes have
cus was on individual freshwater ecosystems, provid- milder winters with temperatures around 13°C. Sum-
ing insights into taxonomic richness, environmental mers are characterized by hot and dry conditions, with
influences and life history strategies. Despite the nar- temperatures ranging from 30 to 39°C (Dambri et al.,
row focus on isolated habitats, these studies provided 2020). Sampling in this region included four temporary
valuable insights into complex ecological dynamics. streams (Figure 2c) and one sebkha.
The culmination of these efforts made a modest con- Finally, the Khenchela region occupies the north-
tribution to substantial knowledge of odonate inter- ernmost tip of the Aurès massif and is located approxi-
actions in North African wetlands, as documented in mately 120 kilometers southeast of Constantine. The
various studies (Bouhala et al., 2019 and references region has different altitudes, ranging from 26 m be-
therein). low sea level (Chott Melghir) to 2169 m above sea level
This study focuses on the relationships between en- (Djebel Chelia). Sampling in the region included seven
vironmental factors (land use and cover, altitude, flow intermittent streams (Figure 2b).

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Sampling use and cover were categorized as “forest”, “rural” (to

describe a forested landscape with scattered dwellings,
We initially selected 76 different stations to cover a formerly referred to as “semi-urban” [Boucenna et al.,
range of aquatic ecosystems, including streams, riv- 2023]), “agriculture” and “pastoralism” (grazing). In
ers (wadis), lakes (garaets) and freshwater reservoirs. addition, in situ measurements were taken for critical
These stations were selected to cover a wide range of environmental parameters such as flow velocity, water
altitudes, ranging from 14 m above sea level in low- depth and bed width at each sampling site. Substrate
lying areas to a maximum altitude of 1608 m. In ad- composition was assessed using established categories
dition, the selected sites were distributed across dif- adopted from Tachet et al. (2010). Hydroperiods (du-
ferent land use systems and climatic zones. Sampling ration of inundation) were dichotomously classified as
was conducted monthly at 40 selected sites (Table 1) permanent or intermittent. Helophyte and hydrophyte
over a twelve-month period spanning from June 2019 cover were determined by visual inspection and ex-
to May 2021. It is worth noting that there was one in- pressed as a percentage. The comprehensive details of
terruption in sampling due to COVID-19 confinement these measurements provide a snapshot of the envi-
measures (April and May were sampled in 2021). GPS ronmental characteristics at the sampled sites (Table 2).
coordinates and altitude were noted for each station. At each site, adult odonates were collected by aerial
Eleven environmental variables (land use, altitude, net while slowly and repeatedly walking a 50 m transect
current velocity, density of helophytes, density of hy- for a total of 20 minutes. In addition, larvae were sys-
drophytes, bed width, substrate, hydroperiod, water tematically collected from different microhabitats using
depth, longitude and latitude) were considered. Land dip nets. Systematic kick sampling was carried out by

Figure 1. Map of north-eastern Algeria with location of sampling sites.

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Table 1. List of sampled sites at the study area. Alt: Altitude.

Latitude Longitude Alt Hydro- Latitude Longitude Alt Hydro-

Code Name Code Name
(N) (E) (m) period (N) (E) (m) period

L1 O. Beni Zid 36.9071889 6.4756389 79 Permanent L23 Jemott_2 35.6510694 7.0118417 829,2 Temporary
L2 O. Tizagbane 37.0109472 6.3748806 492 Permanent L24 Ourkis Dam_1 35.8969056 6.9274639 946,5 Permanent
L3 O. Tamanart_S1 37.0305278 6.5054389 47 Permanent L25 Ourkis Dam_2 35.8930000 6.9300667 946,7 Permanent
L4 O. Tamanart_S2 37.0489000 6.5147083 14 Permanent L26 Touzline_1 35.9376028 6.9373528 994,8 Permanent
L5 O. Kfayoun 36.9378333 6.3020306 194 Temporary L27 Touzline_2 35.9383417 6.9388472 996,6 Permanent
L6 O. Zhor_S1 36.9070333 6.3196333 28 Permanent L29 O. Zerhaib_0 35.8511000 6.3823444 798,4 Temporary
L7 O. Zhor_S2 36.9142611 6.2914694 18 Permanent L33 Tinsilt_2 35.8992306 6.4768556 785,9 Temporaire
L9 O. Guebli_S1 36.8008000 6.6651361 61 Permanent L37 L.Ezzemoul_1 35.8890083 6.5061778 791,2 Temporaire
L10 O. Guebli_S2 36.8600722 6.6151278 31 Permanent L44 O. Lazrag_1 35.3587528 6.7512194 1214 Permanent
L11 O. Gratam 36.7937250 6.6266528 108 Temporary L45 O. Lazrag_2 35.3579417 6.7536528 1201 Permanent
L13 O. Dhimine 35.9969889 7.2797194 803,2 Temporary L51 O. Ibikan_2 35.4056667 6.9662167 1110 Temporary
L14 O. Dahmen_0 36.0086889 7.2337028 817,7 Permanent L52 O. Ibikan_3 35.4120000 6.9641500 1071 Permanent
L15 O. Dahmen_1 36.0203333 7.2487500 799,3 Permanent L53 O. Ibikan_4 35.4185167 6.9502333 1038 Permanent
L16 O. Dahmen_2 36.0406778 7.2732306 779 Permanent L63 O. Rbaa_S2 35.4580333 6.5185667 1040 Permanent
L17 O. Dahmen_3 36.0533583 7.3260972 754,5 Permanent L64 O. Rbaa_S3 35.4898333 6.5354500 1011 Permanent
L18 O. Dahmen_4 36.0581139 7.3401889 746,9 Permanent L66 O. Lakhal_S2 35.3686833 6.6508500 1303 Temporary
L19 Timerganine_1 35.6486528 6.9646778 836 Temporary L67 O. Lakhal_S3 35.3807667 6.6504167 1246 Temporary
L20 Timerganine_2 35.6469194 6.9647889 835,3 Temporary L68 O. Bouilef_S1 35.6173917 6.1872972 1033 Permanent
L21 Timerganine_3 35.6450000 6.9658139 836,1 Temporary L70 O. Bouilef_S3 35.5971722 6.2145861 1000 Temporary
L22 jemott_1 35.6455667 7.0125472 832,7 Temporary L73 O. Labiod_S3 35.0504333 6.1682833 1226 Temporary

Figure 2. Four views of sampling sites: (a) O. Guebli_S1 (L9, Collo); (b) O. Ibikan_S3 (L52, Khenchela); O. Bouzina_S2 (L75,
Batna), Touzline_1 (L26, Oum El Bouaghi).

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Table 2. Physical characteristics of sampled localities.

Helophyte Hydrophyte
Current Bed width Water depth
Code vegetation vegetation Substrate Land use
velocity (m/s) (m) (cm)
(%) (%)
L1 0.47 ± 0.21 4.33 ± 0.58 16.64 ± 2.2 0 0 Sand, Pebbles-boulders Forest
L2 0.47 ± 0.12 5.6 ± 0.75 21.33 ± 1.72 3 0 Sand, Pebbles-boulders Forest
L3 0.44 ± 0.25 6.68 ± 0.95 19.83 ± 5.37 0 0 Sand, Pebbles-boulders Forest
L4 0.58 ± 0.29 8.83 ± 2.82 20.83 ± 5.72 1 1 Sand, Pebbles-boulders Forest
L5 0.26 ± 0.22 1.64 ± 0.67 6 ± 5.24 3 0 Sand, Pebbles-boulders Forest
L6 0.64 ± 0.18 13.12 ± 2.57 27.33 ± 10.59 1 0 Sand, Pebbles-boulders Forest
L7 0.56 ± 0.15 8.7 ± 1.42 39.63 ± 14.66 15 0 Sand Agriculture
L9 0.65 ± 0.45 7.58 ± 1.32 14.67 ± 4.27 15 0 Sand Rural
L10 0.71 ± 0.23 11.73 ± 0.42 34.33 ± 4.89 7 0 Sand Rural
L11 0.11 ± 0.01 1 ± 0.7 2.5 ± 0.7 0 0 Sand, Pebbles-boulders Rural
L13 0.17 ± 0.22 1.5 ± 0.42 24 ± 3.28 5 2 Silt, Pebbles_boulders Agriculture
L14 0.05 ± 0.04 1 ± 0.34 16.63 ± 2.5 10 1 Sand, Pebbles_boulders Agriculture
L15 0.06 ± 0.04 2 ± 1.44 20.5 ± 4.5 3 1 Sand, Pebbles_boulders Agriculture
L16 0.06 ± 0.05 1.66 ± 0.45 28.3 ± 3.23 3 5 Sand, Pebbles_boulders Agriculture
L17 0.09 ± 0.03 8.19 ± 1.15 27.44 ± 5.59 30 1 Silt, Sand Agriculture
L18 0.47 ± 0.22 2.35 ± 0.36 23.16 ± 3.76 5 1 Silt, Sand, Pebbles_boulders Agriculture
L19 – 21.25 ± 5.15 – 40 1 Silt Agriculture
L20 – 23.54 ± 4.89 – 5 2 Silt Agriculture
L21 – 20.38 ± 3.46 – 30 2 Silt Agriculture
L22 – 30.17 ± 8.91 – 6 6 Silt, Sand Pastoralism
L23 – 26.3 ± 6.5 – 5 5 Silt, Sand, Pebbles_boulders Pastoralism
L24 – 31.75 ± 4.59 – 1 1 Silt, Sand, Pebbles_boulders Pastoralism
L25 – 25.38 ± 4.98 – 1 2 Silt, Sand, Pebbles_boulders Pastoralism
L26 – 25.36 ± 6.14 – 5 1 Silt Agriculture
L27 – 34.55 ± 5.65 – 15 2 Silt Agriculture
L29 0.064 ± 0.06 2.13 ± 0.2 14.5 ± 3.27 5 2 Silt Agriculture
L33 – 12.33 ± 3.21 – 5 1 Silt, Sand Agriculture
L37 – 18.88 ± 3.68 – 5 1 Silt Agriculture
L44 2.68 ± 0.43 14.83 ± 0.38 15.08 ± 4.56 5 1 Silt Rural
L45 2.6 ± 7.23 19.83 ± 0.57 34.33 ± 13.73 5 4 Silt Rural
L51 2.06 ± 0.79 9.16 ± 2.88 16.5 ± 8.87 4 0 Clay, Boulders Forest
L52 2.55 ± 0.51 20 ± 0 13.33 ± 3.02 3 0 Silt Rural
L53 2.98 ± 0.61 40 ± 0 13.41 ± 5.99 4 1 Clay Agriculture
L63 0.13 ± 0.04 4.043 ± 1.63 22.69 ± 4.79 20 5 Clay, boulders Pastoralism
L64 0.37 ± 0.3 3.068 ± 1.75 16.15 ± 13.29 50 20 Silt, Pebbles and boulders Rural
L66 0.35 ± 0.24 5.019 ± 1.34 16.85 ± 3.26 30 15 Pebbles, Sand and boulders Agriculture
L67 0.49 ± 0.26 5.062 ± 1.37 25.46 ± 6.29 60 20 Clay, boulders Rural
L68 0.32 ± 0.13 3.032 ± 0.69 22.11 ± 8.11 30 15 Silt, Sand and boulders Forest
L70 0.31 ± 0.11 2.058 ± 0.69 12 ± 3.9 40 10 Clay, Sand and Pebbles Agriculture
L73 0.41 ± 0.37 8.028 ± 2.22 16.22 ± 3.23 10 20 Silt, Pebbles Agriculture

walking slowly and repeatedly over all microhabitats, total abundance was calculated cumulatively for each
covering an area of 100 m² for fifteen minutes. All col- odonate species. Due to the limited sample size of
lected larvae were stored in 80% ethanol until subse- adults, their separate analysis was not considered justi-
quent sorting and identification in the laboratory. fied and the analyses focused solely on the larvae. To
ensure the robustness of our statistical analyses, we
Statistical analysis conducted an exploratory data analysis (EDA) following
the guidelines of Zuur et al. (2010). This preliminary in-
The measurements of environmental variables (flow vestigation aimed to identify potential violations of the
velocity, water depth, helophytes and hydrophytes) underlying assumptions of the statistical techniques
were averaged over the entire study period, while the used. Pairwise Pearson correlations showed strong col-

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

linearity between altitude and latitude. The density of the rich odonate biodiversity in our study area. Most
of helophytes also showed a high correlation with the notably, the families Coenagrionidae, Gomphidae and
density of hydrophytes. In addition, the EDA, through Libellulidae were the most widely distributed. Diver-
graphical plots, indicated potential spatial correlations sity within the Libellulidae stood out with seven spe-
for altitude, flow velocity, bed width, helophytes and cies, followed by the Coenagrionidae with five species.
hydrophytes. Lestidae and Aeshnidae ranked third with three species
Given these results of the EDA, it was decided to each. Calopterygidae, Platycnemididae and Gomphidae
exclude the covariates “latitude” and “hydrophytes” each had a single representative. Two species, Lestes
from further analyses. This step was taken to ensure virens and Orthetrum chrysostigma, were only record-
that subsequent statistical analyses were not unduly ed in their adult stage, while Erythromma viridulum,
influenced by potentially collinear or redundant vari- Boyeria irene and Onychogomphus uncatus were only
ables, thereby improving the interpretability and reli- recorded in their larval stage.
ability of our final results. A non-metric multidimen-
sional scaling analysis (NMDS) based on Bray-Curtis Abundance
dissimilarity was performed on a matrix of 37 sites
and 18 species to find a low-dimensional mapping to Among the recorded species, O. uncatus and Ischnura
Euclidean space that best preserves the rank order of graellsii were the most abundant (Figure 3a). The Tizag­
the dissimilarity matrix. To account for asymmetry in bane site (L2, Collo) was characterized by exceptional
the distribution of species, a Hellinger transformation, abundance of odonates, in contrast to other streams
known for its usefulness in pre-processing abundance in the same vicinity, which had low populations (Fig-
data (Legendre & Borcard, 2018), was applied to the ure 3b). Interestingly, selected wetlands in the Oum El
relative abundances of species before performing a Bouaghi region (L19, L21-23, L26) showed a remarkable
two-dimensional NMDS. abundance of odonates, a phenomenon not reflected
Environmental vector fitting using linear regression in the streams of the Batna region, characterized by
was performed using the “envfit” function to assess lower abundance.
the correlation between the environmental variables
(both continuous variables and centroids of the levels Species richness
of class variables) and the sample ordination. Similarly,
the function “ordisurf” was used to fit a smooth sur- A closer look at species distribution revealed that
face using a thin plate spline fit in generalized addi- I. graellsii and Anax parthenope were the most wide-
tive models (GAM) and interpolate the fitted values. spread, spanning 16 sites (Table 3). However, this dis-
To confirm the influence of significant environmental tribution pattern did not extend uniformly to the two
factors on the odonate assemblages, a permutation Maghrebian endemics, Platycnemis subdilatata (LC)
analysis of variance (PERMANOVA) was performed us- and Enallagma deserti (LC). While P. subdilatata was
ing the function “adonis2” with 1000 permutations. widespread, E. deserti remained restricted to specific
The “betasurf” function was used to assess significant sites in the Hauts Plateaux. The range of species rich-
spatial variation in the dispersion of multivariate eco- ness, which ranged from one to nine, reflected the pat-
logical data. This analysis was particularly important in tern of odonate abundance. Lotic and lentic habitats in
cases where conventional PERMANOVA had difficulty the Oum El Bouaghi region proved to be the hotspots
distinguishing between location and dispersion effects of species richness, while certain wadis in the Collo and
(Anderson et al., 2006). To assess the importance of Batna regions had much lower values (Figure 4).
fitted models and to preserve the spatial dependence
between stations, we used restricted permutation Non-Metric Dimensional Scaling (NMDS)
tests where the data were shuffled with completely
randomization within blocks (regions). NMDS, vector The two-dimensional mapping by NMDS converged to
fitting and restricted PERMANOVA tests were per- a stress value of 0.06 and an R² value between the or-
formed using the ‘vegan’ package (Oksanen et al., dination distance and observed (Euclidian) similarity of
2022) in the statistical software R (R Development 0.99. The NMDS map indicated relatively distinct odo-
Core Team, 2023). nate assemblages (Figure 5) with close similarities be-
tween Collo and Khenchela, both well separated from
Oum El Bouaghi and Batna. The lentic and resource-rich
Results sites (L22, L23, L26, L27) in the Oum El Bouaghi region,
characterized by an assemblage of odonates made up
We documented 2136 individuals, including 1948 lar- of Sympecma fusca, Sympetrum fonscolombii, S. strio-
vae and 188 adults, at 40 different sites. The results of latum and Crocothemis erythraea, was separated from
our study reveal a diverse odonate group comprising 21 the rest.
species distributed in seven families. Of particular note, In addition, the NMDS ordination effectively sepa-
our results include ten Zygoptera and eleven Aniso­ rated the lotic habitats of Oum El Bouaghi and, to a
ptera species (Table 3), providing a detailed snapshot lesser extent, Batna, from their counterparts in Collo

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Table 3. Checklist of odonates recorded in the study area with list of localities.

N Family Species Locality Region

Calopterygidae
1 Calopteryx haemorrhoidalis (Vander Linden, L1, L2, L3, L4, L5, L18, L44, L45, L51, Collo, Oum El
1825) L53, L70 Bouaghi, Aures
Lestidae
2 Lestes virens (Charpentier, 1825) L22 Oum El Bouaghi
3 Chalcolestes viridis (Vander Linden, 1825) L11; L14, L16, L18, L66, L67 Collo; Oum El
Bouaghi, Khenchela
4 Sympecma fusca (Vander Linden, 1820) L22, L23, L26, L27, L33 Oum El Bouaghi
Coenagrionidae
5 Enallagma deserti (Selys, 1871) L21, L24, L26 Oum El Bouaghi
6 Ischnura graellsii (Rambur, 1842) L7, L13, L14, L15, L16, L17, L19, L20, Collo; Oum El
L21, L22, L23, L24, L25, L26, L27, L37 Bouaghi
7 Coenagrion mercuriale (Charpentier, 1840) L13, L15, L16, L17 Oum El Bouaghi
8 Coenagrion caerulescens (Fonscolombe, 1838) L13, L17, L18 Oum El Bouaghi
9 Erythromma lindenii (Selys, 1840) L25
Platycnemidae
10 Platycnemis subdilatata Selys, 1849 L7, L13, L14, L15, L16, L17, L18, L44, Collo, Oum El
L45, L63, L64 Bouaghi, Aures
Aeshnidae
11 Anax imperator Leach, 1815 L37 Oum El Bouaghi
12 Anax parthenope (Selys, 1839) L13, L14, L15, L16, L17, L18, L19, L20, Oum El Bouaghi
L21, L22, L23, L25, L26, L27, L33, L37
13 Boyeria irene (Fonscolombe, 1838) L1, L2, L3, L5, L7, L67 Collo, Khenchela
Gomphidae
14 Onychogomphus uncatus (Charpentier, 1840) L1, L2, L3, L4, L5, L7, L9, L10, L44, L45, Collo, Khenchela
L51, L52, L53, L66
Libellulidae
15 Orthetrum nitidinerve (Selys, 1841) L13, L14, L15, L16, L17, L18, L20, L21, Oum El Bouaghi
L22, L23, L24, L26, L27
16 Orthetrum chrysostigma (Burmeister, 1839) L14, L68 Oum El Bouaghi,
Aures
17 Orthetrum coerulescens (Fabricius, 1798) L6, L44, L45 Collo, Khenchela
18 Sympetrum fonscolombii (Selys, 1840) L14, L16, L19, L20, L21, L22, L23, L24, Oum El Bouaghi
L26, L27, L33, L37
19 Sympetrum striolatum (Charpentier, 1840) L14, L15, L16, L21, L22, L23, L27, L29, Oum El Bouaghi
L33
20 Crocothemis erythraea (Brullé, 1832) L22, L26, L27 Oum El Bouaghi
21 Trithemis annulata (Palisot de Beauvois, 1807) L6, L15, L16 Collo, Oum El
Bouaghi

and Khenchela. Odonate assemblages represented by Vector fitting of environmental variables

such species as Chalcolestes viridis, Platycnemis sub-
dilatata, Coenagrion caerulescens, C. mercuriale and The function “envfit” allowed overlaying of measured
Orthetrum nitidinerve thrived in the relatively open environmental vectors to odonate assemblages. Results
wadis of Oum El Bouaghi and Batna. In contrast, Calo­ of 1000 permutations indicated that seven out of nine
pteryx haemorrhoidalis, Onychogomphus uncatus, variables (land use, altitude, current velocity, helophytes,
Boyeria irene and Orthetrum coerulescens showed a substrate, bed width, water depth, longitude and hydro-
preference for wadis at mid- to high altitudes, often period) were significantly related to the NMDS ordina-
surrounded by forest galleries. A small group of odo- tion differentiating the four regions. Current velocity (R² =
nates, including I. graellsii, A. parthenope, and Trithe- 0.40, p = 0.001), land use (R² = 0.59, p = 0.001), altitude
mis annulata, exhibit high adaptability by frequenting (R² = 0.26, p = 0.005), helophytes (R² = 0.37, p = 0.001),
both lotic and lentic habitats, especially in the Oum El substrate (R² = 0.36, p = 0.003), bed width (R² = 0.27, p =
Bouaghi region. 0.008), and hydroperiod (R² = 0.15, p = 0.021) were lin-

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Figure 3. Species abundance in the study area (a); abundance of odonates in sampled localities (b).

International Journal of Odonatology │ Volume 27 │ pp. 172–186 179

Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Figure 4. Species richness of odonates at sampled localities.

Figure 5. NMDS Bray-Curtis ordination of odonate assemblages from four regions of north-eastern Algeria.

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

early correlated with the NMDS ordination, whereas lon- + Helophytes + Substrate+ Hydroperiod + Longitude), a
gitude and water depth were not (p > 0.05). In addition, restricted PERMANOVA analysis of odonate assembag-
the use of the function of ordisurf indicated a significant es of the study area suggested that only bed width (R² =
nonlinear relationship of longitude with the NMDS ordi- 0.06; p = 0.01) (Figure 6), current velocity (R² = 0.08;
nation (p < 1.9e-05; deviance explained = 61%). p = 0.03), and to a lesser extent, land use (R² = 0.23; p =
To assess the importance of the fitted model (Od 0.06) and altitude (R² = 0.03; p = 0.06) were significant
~ Land_use + Bed_width + Altitude + Current_Velocity (Figure 7).

Figure 6. Relationships between odonate assemblages and bed width (m).

Figure 7. Relationships between odonate assemblages and measured environmental factors: Flow velocity (ms-1): color gradi-
ent, land use: symbols, and latitude (m a.s.l.): size.

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

Discussion velocity, land cover and use, and altitude. Although

riverbed width is known to influence odonate assem-
A total of 21 species were documented in this study, blages in lotic environments (Kietzka et al., 2014), this
including ten Zygoptera and eleven Anisoptera. This factor is collinear with habitat type in our study, con-
represents a significant proportion – about 33% of the firming the different ecological requirements between
total species inventory recorded in Algeria (Samraoui & lotic and lentic odonates.
Menaï, 1999) and 47% of Numidian species (Samraoui Another significant environmental factor, flow veloc-
& Corbet, 2000). As was found in the main tributary of ity, shows remarkable correlations with odonate as-
the River Seybouse (Bouhala et al., 2019), the Ibero- semblages, as also found in other studies (Bouhala et
Maghrebian I. graellsii and the Maghrebian P. subdilata­ al., 2019; Hofmann & Mason, 2005; Sato & Riddiford,
ta were among the most abundant and widespread 2008). Flow velocity is closely related to the dissolved
species. Noteworthy species missing in this survey in- oxygen content of the water and it underscores its
clude the Maghrebian endemics Calopteryx exul Selys, role in shaping distribution patterns (He et al., 2011).
1853 (EN) and Gomphus lucasii Selys, 1849 (EN), con- The influence of oxygen on odonate larvae is crucial,
sidered threatened on the Mediterranean and North as oxygen levels can affect their respiration, behaviour
African IUCN Red Lists (Riservato et al., 2009; Samraoui and adaptation to different environments (Apodaca &
et al., 2010). Chapman, 2004). This, in turn, influences distribution
The absence of larvae in the two species recorded as patterns and emphasises the crucial role that hydrody-
adults, Lestes virens and Orthetrum chrysostigma, can namic conditions play in shaping the suitability of habi-
be attributed to the short duration of their larval de- tats for particular species. A deeper understanding of
velopment. On the other hand, the absence of adults these relationships improves our knowledge of habitat
in Boyeria irene and Onychogomphus uncatus could be preferences and the potential impacts of anthropogen-
due to their behaviour. The foraging activity of B. irene ic changes on these habitats.
usually peaks around sunset, while the appearance of Altitude was found to be marginally significant. This
O. uncatus at their breeding sites can be sporadic. Final- may be due to the fact that this factor may interact with
ly, the absence of adults of E. lindenii could be due to a the local climate, as may happen in the Aurès Moun-
small breeding population. tains (Batna and Khenchela), highlighting the differenti-
The complex patterns of species abundance and dis- ated character of habitat conditions. Altitude has been
tribution showed a remarkable correlation with habi- found to be a key environmental factor in driving odo-
tat morphology, hydrology and landscape cover. These nate assemblages in numerous studies (Campbell et al.,
environmental aspects, recognized as major drivers of 2010; Gomez-Anaya et al., 2011; Lee et al., 2018; Sam-
diversity, are particularly true for macroinvertebrates ways, 1989). Similarly, land use was observed to have
(Resh et al., 2002; Rocha-Ortega et al., 2019). As a key a marginal but nevertheless recognisable influence on
indicator of biodiversity, species richness plays a cen- shaping odonate assemblages in north-eastern Alge-
tral role in assessing the overall health of ecosystems ria. This result is in line with a number of studies that
(Magurran, 1988). Previous studies on North African emphasise the importance of landscape structure for
rivers have shown a positive correlation between spe- odonate distribution (Le Gall et al., 2018; Samways &
cies richness and riverine vegetation cover (Bouche- Steytler, 1996; Smith et al., 2022). These studies high-
louche et al., 2015). Changes in land use, combined light the complex interplay between land use practices
with the overarching influence of global warming, have and odonate community composition and provide valu-
been shown to lead to significant shifts in the species able insights into the factors influencing odonate diver-
composition of odonate assemblages (Morghad et al., sity in this region.
2019). The observed direct relationship between spe- The responses of odonate assemblages to various
cies richness and habitat heterogeneity, particularly in environmental changes have attracted much atten-
the Oum El Bouaghi region, highlights the habitat-spe- tion in the ecological literature (Corbet, 1999). Signifi-
cific preferences and ecological tolerances of different cant factors influencing these responses are shade or
odonate species (Deacon et al., 2019). This latter area, light exposure (Remsburg et al., 2008; Worthen et al.,
a drought-prone area, hosted early colonizers of new- 2021) and substrate characteristics (Henrikson, 1993;
ly formed habitats such as I. graellsii, Anax imperator, Marques Pires et al., 2020). A key biotic factor influenc-
Sympetrum fonscolombii, S. striolatum and Trithemis ing odonate distribution is the presence of vegetation,
annulata (Márquez-Rodríguez et al., 2023; Morghad et which is important in both lotic habitats (Barbosa de
al., 2019). Oliveira-Junior et al., 2017; Buchwald, 1992) and lentic
The significant correlations observed between odo- environments, whether in natural contexts (Raebel et
nate assemblages and specific environmental variables al., 2011) or in urban landscapes (Goertzen & Suhling,
provide invaluable insight into the factors that deter- 2012).
mine their distribution. The regional shifts in odonate Contrary to expectations, another notable environ-
assemblages in northeastern Algeria are best explained mental factor that did not appear to have a significant
by their associations with a set of key environmental influence on odonate composition in our study is the
variables that include habitat type (lentic vs lotic), flow hydroperiod. The hydroperiod gradient, along with

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Satour, Hezil, Taferghoust ... Samraoui Unraveling environmental determinants of odonate assemblages in northeastern Algeria

physical and biotic factors, plays a crucial role in shaping regions would have undoubtedly improved the study
community structure (Wellborn et al., 1996). Although which did not include the shallow freshwater lakes and
there is a considerable overlap in species composition brackish marshes of Numidia. Similarly, the study area
between intermittent and perennial streams (del Rosa- lacked large flood plain rivers. Expanding the study to
rio & Resh, 2000), taxa with shorter life cycles tend to exuviae and larvae in addition to adults could also pro-
dominate habitats with intermediate to short hydrope- vide a more comprehensive view of assemblage struc-
riods (Marques Pires et al., 2017a, b). tures in these wadis. Consideration of these limitations
This emphasises the importance of temporary, fish- directs future research in promising directions, such as
less habitats that develop during dry periods and pro- the study of interactions between specific environmen-
vide a unique opportunity for odonates to thrive (Dorn, tal factors and odonate life histories. It is expected that
2008; Wellborn et al., 1996). Odonate communities a thorough investigation of these interactions could re-
in intermittent habitats have species with specialised veal hidden ecological relationships and thus improve
adaptations that allow them to withstand the hot, dry our understanding of odonate assemblages.
Mediterranean summers (Aguero-Pelegrin & Ferreras-
Romero, 1992; Samraoui, 2009; Samraoui et al., 1998). Acknowledgments
The temporal rhythm inherent in Mediterranean aquat-
ic systems remains of paramount importance and em- We are most grateful to two anonymous reviewers for their comments
phasises the ecological dynamics that determine the and helpful suggestions. This study was supported by the Algerian
Ministère de l’Enseignement Supérieur et de la Recherche Scientifique
distribution of odonates and their responses to envi-
(M.E.S.R.S./P.R.F.U.).
ronmental gradients.
All of these studies expanded our understanding of
odonate responses to environmental change and shed References
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