Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 August 2026 | 18(8): 29375–29389
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.10551.18.8.29375-29389
#10551 | Received 24 March 2026 | Final received 26 June 2026| Finally
accepted 22 July 2026
Bat
assemblages at two contrasting agricultural sites in northern Algeria: a
preliminary conservation assessment
Saida Meghezzi
1 , Fatima Zohra Bissaad 2 , Farid Bounaceur 3 , Afaf Boukhari
4 ,
Moncef Ait-Abdesselam
5 & Leila Benfekih 6
1,6 Laboratory for Research on
Medicinal and Aromatic Plants, Department of Biotechnology and Agroecology,
Faculty of Nature and Life
Sciences, Université of Blida 1, Route de Soumaa, Blida 09000, Algeria.
2 Laboratoire de Recherche de Bioinformatique, Microbiologie Appliquée et Biomolécules
(LBMAB), Faculté des Sciences,
Université M’Hamed
Bougara, Boumerdes 35000,
Algeria.
3 Équipe de Recherche Biologie
de la Conservation en Zones Arides
et Semi Arides, Laboratoire
Agronomie Environnement, Institut des Sciences, Université
de Tissemsilt, Tissemsilt
38000, Algeria.
4 Laboratoire Technologies Douces,
Valorisation, Physico-Chimie
des Matériaux Biologiques
et Biodiversité, Université
M’Hamed Bougara, Boumerdes 35000, Algeria.
5 Laboratoire d’Hygiène et Pathologie Animales, Faculté des Sciences de
la Nature et de la Vie, Université Ibn Khaldoun, Tiaret 14000, Algeria.
1 saida.meghezzi@yahoo.com, 2 f.bissaad@univ-boumerdes.dz
(corresponding author), 3 farid.bounaceur@univ-tissemsilt.dz, 4
afafboukhari0@gmail.com, 5 moncef.agrovet@gmail.com, 6 leila.benfekih@univ-blida.dz
Editor: Bhargavi Srinivasulu,
Zoo Outreach Organisation, Hyderabad, India. Date of publication: 26 August
2026 (online & print)
Citation: Meghezzi, S., F.Z.
Bissaad, F. Bounaceur, A. Boukhari, M. Ait-Abdesselam &
L. Benfekih (2026). Bat assemblages at two
contrasting agricultural sites in northern Algeria: a preliminary conservation
assessment. Journal of Threatened Taxa 18(8): 29375–29389. https://doi.org/10.11609/jott.10551.18.8.29375-29389
Copyright: © Meghezzi et al. 2026. Creative Commons Attribution 4.0
International License. JoTT allows unrestricted use,
reproduction, and distribution of this article in any medium by providing adequate
credit to the author(s) and the source of publication.
Funding: No specific funding was received for this study.
Competing interests: The authors declare no competing interests.
Author details & Author contributions: See end of this article.
Acknowledgements: The authors sincerely thank M. Taharout Djamel, the farmer, for kindly granting access to his agricultural property and allowing us to conduct this study on his farms.
Abstract: Agricultural intensification is
a key driver of biodiversity loss in Mediterranean agroecosystems, yet its
effects on protected bat assemblages remain poorly documented in northern
Africa. All six bat taxa recorded are legally protected in Algeria. Several of the
recorded species are also covered by international conservation instruments for
their European populations, while the probable North African endemic Plecotus cf. gaisleri
is of particular biogeographic interest. We conducted a preliminary assessment
of bat activity, diversity, and community composition at two representative
agricultural sites—a low-input citrus orchard and an intensively managed
vineyard—in northern Algeria. Acoustic monitoring was conducted from March to
November 2023. A total of 349 acoustic contacts were recorded at the citrus
orchard site and 74 at the vineyard site, corresponding to a 4.6-fold
difference in observed activity. Negative binomial modelling indicated a
substantially lower fitted contact rate at the vineyard than at the orchard
(IRR = 0.215, 95% CI: 0.130–0.355). Monthly taxon richness and Shannon
diversity were also higher at the orchard. The vineyard assemblage was strongly
dominated by Pipistrellus kuhlii, whereas a more taxonomically diverse assemblage
was detected at the orchard. NMDS suggested site-associated structuring of bat
assemblages, and a paired PERMANOVA detected a significant difference between
the two sampled sites (R² = 0.281, pseudo-F = 6.26, p = 0.004).
No statistically significant difference in multivariate dispersion was detected
(PERMDISP: F = 0.808, p = 0.100), although the small sample size
limits the power of this diagnostic test. Although based on a single
orchard–vineyard pair, these results suggest that the low-input orchard
examined here may provide favourable conditions for a
protected bat assemblage. Replicated multi-site studies are needed before
generalizing this pattern to low-input perennial agroecosystems across North
African farmland.
Keywords: Acoustic monitoring,
agroecosystems, biodiversity conservation, community composition, low-input
farming, Mediterranean landscapes, preliminary assessment.
INTRODUCTION
Agricultural pests are a major constraint on crop productivity
and represent a significant source of economic loss in global agroecosystems (Oerke 2006). In Algeria, pest-driven reductions in
agricultural output intersect with broader economic vulnerabilities linked to
hydrocarbon dependency and climate variability, increasing the need for
resilient and sustainable agricultural systems (Bouchenak
2021). Climate change is expected to intensify pest pressure through shifts in
phenology, distribution, and trophic interactions (Deutsch et al. 2018; Lehmann
et al. 2020), further reinforcing the urgency of effective pest management
strategies.
Integrated Pest Management (IPM) provides a coherent
framework for reducing reliance on synthetic pesticides while maintaining crop
yields and profitability (Pretty & Bharucha 2014).
IPM combines biological control, habitat management, resistant cultivars,
diversified cropping systems, and decision-support tools to prevent or limit
pest outbreaks (Roubos et al. 2014; Gurr et al. 2017). Beyond direct pest suppression, IPM can
enhance biodiversity and long-term agroecosystem resilience by promoting
functional diversity and ecosystem services (Gupta et al. 2024; Marcelino et
al. 2024). These approaches are particularly relevant in Mediterranean
agricultural systems, where pest outbreaks and pesticide resistance generate
substantial economic and environmental costs (Biondi et al. 2012).
Insectivorous bats constitute a potentially important,
yet underutilized, component of biological control in agricultural landscapes.
Numerous studies show that bats consume large proportions of agricultural pest
taxa, often exceeding 40% of their diet in intensive crop systems (Zhu et al.
2024), and bats have been estimated to provide agricultural pest-control
services worth at least USD 3.7 billion annually in North America, although
such estimates remain context-dependent (Boyles et al. 2011; Maas et al. 2016).
In Mediterranean contexts, bat assemblages exploit agricultural pest prey and
can provide ecologically and economically relevant pest regulation services
(Russo & Ancillotto 2015; Ancillotto
et al. 2022). The efficiency of these ecosystem services depends on bat species
composition, habitat structure, landscape complexity, and exposure to
agrochemical inputs (Tuneu-Corral et al. 2023).
Structurally complex agricultural habitats with permanent vegetation generally
support greater insect abundance and foraging opportunities, whereas intensive
management and frequent pesticide applications may reduce prey availability and
limit bat activity, thereby decreasing the potential for natural pest
suppression.
In northern Africa, chiropteran research has
predominantly focused on natural, cavernicolous,
urban, and rural habitats, documenting species distributions, roosting ecology,
trophic patterns, and conservation status (Bendjeddou
et al. 2014, 2017, 2020; Farfar et al. 2017; Loumassine et al. 2017a,b, 2019, 2020; Mokrani
et al. 2018a,b). More recently, studies in Algeria have expanded to forest
ecosystems, including Mediterranean montane habitats (Hamida et al. 2021; Ait-Abdesselam et al. 2025; Boukhari
et al. 2025). Collectively, this literature highlights the taxonomic and
functional diversity of Algerian bats across non-agricultural environments.
However, agroecosystems remain largely unstudied, despite being characterized
by high pest pressure, frequent pesticide use, and landscape simplification,
and despite the well-established role of bats as biological control agents in
agricultural systems elsewhere.
Bat taxa recorded in the present study are legally protected
under Algerian national legislation. Several of the identified species are also
covered by international conservation instruments for their European
populations, although these listings should not automatically be interpreted as
applying to the Algerian populations recorded here. Despite their national
legal protection, bat diversity, activity, and habitat use in Maghreb
agroecosystems remain poorly documented, even though agricultural landscapes
occupy extensive areas of the Mediterranean lowlands. Assessing bat assemblages
in agricultural landscapes can therefore provide useful baseline information
for regional conservation planning and the development of biodiversity-friendly
farming practices.
The present study addresses this knowledge gap by characterizing
bat assemblages at two contrasting agricultural sites in the Sebaou Valley, near Draâ Ben Khedda, Tizi Ouzou
Province, northern Algeria. Because only one citrus orchard and one vineyard
were surveyed, the study should be interpreted as a preliminary
conservation-oriented case study rather than as a spatially replicated
comparison of crop types or management systems. Given the preliminary two-site
design, we evaluated three site-specific predictions: (1) monthly taxon
richness and Shannon diversity would be higher at the sampled citrus orchard
than at the sampled vineyard; (2) the vineyard assemblage would show stronger
numerical dominance by generalist taxa, particularly Pipistrellus
spp.; and (3) assemblage composition would differ between the two sampled
sites. Potential relationships with vegetation structure, management intensity,
pesticide use, and prey availability were treated as hypotheses for future
replicated studies rather than as mechanisms directly tested here. By
integrating acoustic activity, α-diversity indices, and community composition
analyses, this study provides a first preliminary assessment of bat communities
in Algerian agroecosystems and highlights their potential conservation value
within Mediterranean agricultural landscapes.
The study was conducted in the Sebaou
Valley, near Draâ Ben Khedda,
Tizi Ouzou Province,
northern Algeria (36.733° N, 3.950° E), at an elevation of approximately 56 m. The area
lies within the sub-humid Mediterranean bioclimatic zone and is characterized
by mild winters. Mean annual temperature is approximately 19.2°C, and annual
precipitation exceeds 600 mm. Soils at both study sites were classified as
sandy clay loam. The surrounding area is predominantly agricultural and
includes several crop types (Image 1).
The
first selected citrus orchard (Site A) is embedded within an extensive
agricultural landscape dominated by long-established citrus orchards,
interspersed with vegetable cropping systems. This orchard is managed under an
extensive conventional production system, characterized by a low planting
density (800–850 trees ha⁻¹), infrequent application of
agricultural inputs, and the absence of weed management practices. Over the
monitoring period spanning March–November 2023, no pesticide treatments were
applied by the farmer (Table 1).
By
contrast, the second selected vineyard is located in an area characterized by
high agricultural production intensity. This table grape vineyard operates
under a high-input intensive production system, featuring a high
planting density (1,111–1,200 vines ha⁻¹), raised-bed cultivation, and drip irrigation combined with
frequent applications of agrochemical inputs, including fertilizers,
herbicides, and pesticides, with the objective of maximizing yield. Weed
management was continuously implemented through the use of chemical herbicides
throughout the study period (Table 1).
Vineyards
in the area are also susceptible to fungal diseases, notably powdery mildew (Uncinula necator)
and downy mildew (Plasmopara viticola), for which farmers applied fungicides such as
Mancozeb and Cyproconazole. To control insect pests, including the African
leafhopper, mites, and grape moths, insecticides such as Deltamethrin and
Chlorpyrifos-methyl were used. Glyphosate was applied for weed control as a
chemical herbicide.
Because
only one citrus orchard and one vineyard were included, these two sites were
considered representative examples of contrasting agricultural management
contexts rather than independent replicates of each habitat type. Therefore,
observed differences in bat activity and assemblage structure should be
interpreted as site-specific patterns that provide preliminary insights for
future multi-site investigations.
Acoustic surveys were conducted monthly from March to
November 2023 at both study sites. Each monthly survey consisted of a one-hour
acoustic monitoring period conducted after sunset during the period of peak bat
activity. The same fixed 500-m transect was traversed four consecutive times,
once during each 15-minute segment (P1–P4), at an approximately constant
walking speed of 2 km h⁻¹. The Batlogger M operated
throughout the monitoring period and stored triggered acoustic files of 5 s at
a sampling frequency of 312.5 kHz. The four 15-minute segments within each
monthly survey were treated as non-independent subsamples and pooled at the
site × month level before statistical analyses. Thus, all inferential analyses
were based on 18 pooled site × month observations. These observations represent
repeated monthly measurements from the same two sites and should not be
interpreted as independent spatial replicates of orchard and vineyard habitats
(Image 2).
At each site, a fixed 500-m transect was traversed four
consecutive times during the one-hour session, with each traversal lasting
approximately 15 min at a walking speed of about 2 km h⁻¹. Successive
traversals were conducted in alternating directions along the same route.
Surveys began at sunset. The two sites were surveyed on consecutive nights,
with each survey beginning at sunset. Survey order was alternated among months:
when Site A was surveyed on the first night and Site B on the following night
during one monthly survey, Site B was surveyed first during the next monthly
survey. Bat calls were recorded using a Batlogger M (Elekon AG, Lucerne, Switzerland; firmware v2.6.2). The
detector operated continuously in automatic trigger mode throughout each
one-hour survey and stored triggered acoustic events as 5-s recordings.
Acoustic signals were sampled at 312.5 kHz, corresponding to a Nyquist
frequency of 156.25 kHz. During each one-hour survey, air temperature and relative
humidity were measured every 15 minutes using an IHM6150SI pocket thermo-hygrometer
(type K) (precision ±1°C; ±5% RH). Cloud cover
and wind speed were also recorded to document
weather conditions potentially influencing bat activity and detectability
(Russo & Jones 2003). Weather variables were recorded to document survey
conditions and to ensure that surveys were not conducted during rainfall or
strong winds. They were not included as model covariates because of the small
sample size, the limited number of observations relative to the number of
potential predictors, and their seasonal collinearity with month. Consequently,
seasonal climatic effects cannot be separated from other sources of
month-related variation.
Recorded calls were manually inspected using BatSound v3.10 (Pettersson Elektronik AB, Sweden). Spectrograms were examined using
call structure, start and end frequencies, frequency of maximum energy,
bandwidth, call duration, and pulse shape. Taxonomic assignments followed Barataud (2012) and regional reference data from
northwestern Africa (Russo & Jones 2003; Disca et
al. 2014; Dalhoumi et al. 2016; Ahmim
et al. 2020).
A total of 1,978 call sequences were recorded during the
sampling period. After manual validation, 423 high-quality sequences (21.39%)
were retained for species identification and acoustic analyses. The remaining
1,555 recordings were excluded (78.61%) because of poor signal quality,
ambiguous acoustic characteristics, or environmental and insect noise.
A bat contact was defined as one triggered 5-s recording
containing a sequence of echolocation calls attributable to a single bat pass.
Recordings containing simultaneous calls from clearly distinguishable taxa were
counted separately for each taxon.
High-duty-cycle calls of Rhinolophus hipposideros
were identified from their characteristic FM/CF/FM structure and a
peak-frequency range of 115.9–121.0 kHz reported for northern Algeria (Ahmim et al. 2020). Because rhinolophid
calls are highly directional and may be detected less readily during walked
transects, only recordings with a clear constant-frequency component and
diagnostic harmonic structure were retained.
Low-duty-cycle calls assigned to Eptesicus isabellinus, Hypsugo savii, Pipistrellus kuhlii, and Pipistrellus pipistrellus were identified from a
combination of FM/QCF structure, frequency of maximum energy, pulse shape, and
regional reference data. Calls falling within known frequency-overlap ranges
and lacking additional diagnostic characteristics were retained only at
acoustic-group level and excluded from species-level analyses.
Calls assigned to Plecotus showed
an FM structure and a starting-frequency range consistent with regional
descriptions of P. gaisleri
(Dalhoumi et al. 2016). However, because species
within Plecotus
cannot be reliably distinguished using acoustic data alone, these records are
reported provisionally as Plecotus cf. gaisleri.
This designation indicates probable, but not definitive, species identity.
The six bat taxa retained for analysis were Rhinolophus hipposideros (Borkhausen,
1797) (Rhinolophidae), Eptesicus isabellinus (Temminck,
1840) (Vespertilionidae), Hypsugo savii (Bonaparte, 1837) (Vespertilionidae), Pipistrellus kuhlii (Kuhl, 1817) (Vespertilionidae), Pipistrellus pipistrellus (Schreber,
1774) (Vespertilionidae), and Plecotus cf. gaisleri (Benda et al., 2004) (Vespertilionidae).
For each sampling session, the dataset contained
species-specific contact counts. Overall bat activity per session was
quantified as the total number of acoustic contacts across
all detected species: Total contacts
=
, where ni is the number
of contacts for species i in that
session. A sampling session was considered positive
when Total contacts > 0. Monthly totals (per site) were obtained by
aggregating the four periods (P1–P4) within each month.
Taxon richness was computed per site × month as the
number of detected taxa (contacts > 0). Assemblage composition was expressed
as the proportion of total contacts contributed by each taxon at each site.
Community diversity was quantified monthly using standard
indices. Shannon diversity was computed as
following Shannon (1948), with
and . Diversity metrics were
computed in R ver
-sion 4.5.2 (R Core Team 2025) following standard community
ecology workflows, including the vegan
package (version 2.7-1) when applicable (Oksanen et
al. 2022).
To compare monthly taxon richness and Shannon diversity
between the citrus orchard and the vineyard, we used the Wilcoxon signed-rank
test for paired comparisons. This non-parametric test was selected because the
paired data did not meet the assumptions of normality. The Wilcoxon signed-rank
test was therefore applied only to taxon richness and Shannon diversity.
Statistical significance was assessed at p < 0.05 using the function wilcox.test in R.
vegan package in R (Oksanen
et al. 2022). The stress value was used to evaluate
ordination quality, with values < 0.1 considered indicative of a good
representation of the data.Site-associated differences in bat assemblage structure
were evaluated using PERMANOVA based on Bray–Curtis dissimilarities,
implemented with the adonis2 function
in the vegan package (Oksanen et al. 2022).
The analysis was conducted on the taxon-by-sample matrix containing the 18
pooled site × month observations. To preserve the paired temporal structure,
permutations of the Site labels were restricted within Month. With nine monthly
pairs, 511 permutations were generated in addition to the observed
configuration. Because only one site represented each agricultural context, the
analysis evaluates a repeated contrast between the sampled orchard and vineyard
and does not provide spatial replication of crop type or management intensity.
Homogeneity of multivariate dispersion was evaluated using betadisper (Anderson 2001) and permutest, with permutations following
the same paired monthly structure.
Monthly bat activity was analysed
using a negative binomial generalized linear model with a log link, implemented
in the glmmTMB package (family = nbinom2;
Brooks et al. 2017). The response variable was the total number of acoustic
contacts recorded during each pooled one-hour site × month session (n = 18).
Site and Month were included as fixed effects, and the fitted model was: Total
contacts ~ Site + Month. Because each agricultural context
was represented by only one site, the Site coefficient is interpreted strictly
as a contrast between the sampled orchard and vineyard rather than as a general
effect of crop type or management intensity. Effect sizes are presented as
incidence rate ratios with 95% confidence intervals obtained by exponentiating
the model coefficients.
Model adequacy was assessed using simulation-based
residual diagnostics implemented in DHARMa
(Hartig 2016), including checks for residual
uniformity, dispersion, and outliers.
All analyses were performed in R (RStudio environment). Figures were produced using ggplot2
(Wickham 2016) and exported at publication-quality resolution.
A total of 423 bat acoustic contacts were recorded across
both agricultural habitats during the sampling period. Bat activity was
substantially higher in the orchard (Site A), with 349 contacts, compared to
the vineyard (Site B), where only 74 contacts were detected, corresponding to a
4.6-fold difference in total activity.
Bat contacts were detected in 33 of the 36 15-min
segments at the orchard site (91.7%) and in 21 of the 36 segments at the
vineyard site (58.3%). Segment-level detection frequencies are reported descriptively
only; all inferential analyses were conducted at the pooled site × month level.
Monthly activity patterns differed markedly between sites (Image 4). In the
orchard, bat activity peaked in March (92 contacts) and remained relatively
high in August (53 contacts) and October (47 contacts), followed by a
pronounced decline in June (4 contacts) and November (18 contacts). In
contrast, vineyard activity remained consistently low throughout the season,
with moderate increases in October (20 contacts) and in July and September (12
contacts each), and minimal activity in June (1 contact) and November (3
contacts).
The negative binomial model indicated a significantly
lower fitted contact rate at the sampled vineyard than at the sampled orchard
(IRR = 0.215, 95% CI = 0.130–0.355, p
< 0.001). The fitted contact rate at the vineyard was approximately 79%
lower than that at the orchard. This estimate represents a contrast between the
two sampled sites and should not be interpreted as a general effect of habitat
type or management intensity. Relative to March, activity was significantly
lower in June and November, whereas the remaining month-specific coefficients
did not differ significantly from the March reference level (Table 3).
Taxon richness was consistently higher in the orchard
than in the vineyard throughout the sampling period (Image 5). A total of six
bat taxa were detected in the orchard, including R. hipposideros,
E. isabellinus, H. savii,
P. kuhlii, P. pipistrellus,
and Plecotus cf. gaisleri. In contrast, only three taxa were recorded in
the vineyard: E. isabellinus, P. kuhlii, and P. pipistrellus.
Monthly taxon richness in the orchard ranged 2–6 taxa,
with a maximum observed in April, whereas vineyard assemblages were frequently
dominated by a single taxon and reached a maximum of three taxa only in August.
Wilcoxon signed-rank tests indicated significantly higher monthly taxon
richness in the orchard (V = 28, p = 0.0213), with median values of
three taxa per month compared to one taxon per month in the vineyard,
corresponding to a median difference of two taxon (Table 4).
Shannon diversity followed a similar pattern (Image 6).
Median Shannon diversity was 0.971 in the orchard and 0 in the vineyard, and
this difference was statistically significant (V = 44, p = 0.0129). Overall, these findings
indicate that the citrus orchard
supported significantly richer and more diverse bat assemblages than the
vineyard.
Taxon composition differed markedly between the two
agricultural habitats (Image 7). In the orchard, bat activity was co-dominated
by P. kuhlii (43.6% of total contacts) and E.
isabellinus (43.3%), with additional
contributions from H. savii (5.7%), P. pipistrellus (4.6%), Plecotus
cf. gaisleri (2.3%), and R. hipposideros (0.6%).
In contrast, the vineyard assemblage was overwhelmingly
dominated by P. kuhlii, which accounted for
93.4% of all contacts, while E. isabellinus
(3.9%) and P. pipistrellus (2.6%) were only
sporadically detected. This overwhelming dominance by a single species was
consistent with the lower taxon richness and Shannon
diversity observed in the vineyard.
Community
structure: NMDS, PERMANOVA, and multivariate dispersion
The NMDS ordination suggested site-associated structuring
of monthly bat assemblages (Image 8). The two-dimensional solution provided a
good representation of the Bray–Curtis dissimilarity matrix (stress = 0.062).
Orchard observations appeared more widely distributed in ordination space than
vineyard observations.
PERMANOVA based on permutations restricted within Month
detected a significant difference in assemblage structure between the two
sampled sites (pseudo-F = 6.26, R² =
0.281, paired permutation p
= 0.004). The repeated contrast between the orchard and vineyard therefore
accounted for 28.1% of the variation in the monthly assemblage data. The test
of multivariate dispersion did not detect a statistically significant
difference between sites (PERMDISP: F
= 0.808, paired permutation p
= 0.100). Thus, there was no statistical evidence that the PERMANOVA result was
explained solely by unequal multivariate dispersion, although the small sample
size limits the power of the dispersion test. Because only one site represented
each agricultural context, these findings describe a repeated difference
between the two sampled sites rather than a general effect of orchard versus
vineyard habitat.
Before discussing the ecological implications of our
findings, an important limitation of the present study should be acknowledged.
Because the study was conducted in only one citrus orchard and one vineyard,
habitat type is confounded with site identity, and the observed differences
cannot be attributed solely to differences in agricultural management
intensity. Consequently, our results should be interpreted as preliminary
evidence highlighting contrasting bat assemblages between two representative
agricultural sites rather than definitive comparisons between cropping systems.
These findings provide a baseline for future replicated, multi-site studies
that incorporate direct measurements of insect prey availability, pesticide
exposure, and other environmental variables to better disentangle the effects
of agricultural management from site-specific characteristics.
Within the limits of this two-site case study, the
sampled citrus orchard consistently showed higher bat activity, richness, and
Shannon diversity than the sampled vineyard. These differences describe
contrasting patterns between the two sites but cannot be attributed
specifically to crop type, management intensity, pesticide use, or vegetation
structure because these factors were not independently replicated.
Several non-exclusive mechanisms could plausibly explain
the higher activity observed at the orchard site. Its perennial canopy,
unmanaged ground vegetation, and lower reported input use may provide greater
structural complexity and potentially more stable prey resources than the
vineyard. However, insect abundance, pesticide residues, surrounding landscape
composition, roost proximity, and microclimatic conditions were not quantified.
These explanations should therefore be treated as hypotheses for future
replicated studies (Russo & Jones 2003; Fahrig et
al. 2011; Park 2015; Azam et al. 2016). Evergreen
orchards, in particular, may sustain insect populations across seasons, thereby
offering more reliable foraging opportunities for bats. Previous studies have
shown that vegetation structure, hedgerows, and landscape heterogeneity can
influence bat activity and prey availability in agricultural landscapes (Boughey et al. 2011; Maas et al. 2016; Froidevaux
et al. 2019). Recent Algerian studies have also documented variation in bat
activity and richness along environmental gradients and among structurally
different habitats (Ait-Abdesselam et al. 2025; Boukhari et al. 2025).
In contrast, the vineyard exhibited low bat activity and
strong dominance of P. kuhlii, a generalist
species widely recognized for its tolerance to habitat simplification and
anthropogenic disturbance (Barataud 2012; Jung &
Threlfall 2016). Such dominance reflects a community simplification process
frequently observed in intensively managed agricultural systems, where habitat
homogenization and chemical inputs act as strong ecological filters (Geiger et
al. 2010; Tscharntke et al. 2012). Vineyards are
often associated with frequent pesticide applications, which can reduce
arthropod abundance and diversity and thereby limit food resources for
insectivorous bats (Wickramasinghe et al. 2004;
Sánchez-Bayo & Wyckhuys
2019). Although pesticide applications were reported at the vineyard site,
neither pesticide exposure nor insect abundance was quantified. The observed
differences therefore cannot be attributed directly to chemical management and
may also reflect vegetation structure, landscape context, roost proximity, or
other site-specific factors.
Although most taxa recorded in this study are currently
classified as Least Concern (LC) globally, and Plecotus cf. gaisleri is not evaluated, global
assessments may not fully reflect regional population trends or local pressures
associated with agricultural intensification. The legal protection of these
taxa in Algeria, together with the limited information available on their use
of northern African agroecosystems, highlights the need for regional baseline
data. Moreover, agri-environment schemes do not
necessarily benefit all bat taxa or biodiversity components in conventional
agricultural landscapes, and some management prescriptions may have limited or
even negative effects (Fuentes-Montemayor et al. 2011; MacDonald et al. 2012;
Park 2015). These considerations underscore the importance of evaluating the
effectiveness of farming practices and conservation measures under local
conditions.
Monthly activity varied over the survey period, with
particularly low values in June and November relative to March. Such variation
may reflect seasonal changes in weather, prey availability, reproduction,
migration, or torpor. Because these mechanisms were not measured directly and
only one annual cycle was sampled, the observed monthly pattern should not be
interpreted as evidence that the orchard buffered seasonal environmental stress
(Russo & Jones 2003; Adams 2010).
Multivariate analyses indicated site-associated
differences in assemblage structure. NMDS suggested partial structuring between
orchard and vineyard observations, and the paired PERMANOVA detected a
significant Site contrast after restricting permutations within Month (pseudo-F
= 6.26, R² = 0.281, p
= 0.004). The test of multivariate dispersion did not detect a statistically
significant difference between sites (PERMDISP: F = 0.808, p
= 0.100). Thus, there was no statistical evidence that the PERMANOVA result was
explained solely by unequal dispersion, although the limited number of monthly
observations reduces the power of this diagnostic test (Anderson 2001; Legendre
& Legendre 2012; Oksanen et al. 2022). These
findings describe contrasting assemblage structures at the two sampled sites
but should not be interpreted as demonstrating a general effect of orchard
versus vineyard habitat because each agricultural context was represented by
only one site.
From a conservation perspective, the higher bat activity
and diversity observed at the sampled orchard suggest that some low-input
perennial agricultural sites may provide favourable conditions for bats. Bats
can provide important ecosystem services in agricultural landscapes,
particularly insect-pest suppression (Boyles et al. 2011; Kunz et al. 2011;
Russo et al. 2018). However, replicated studies are required before this
pattern can be generalized to low-input agricultural systems across the region.
Maintaining vegetation cover and reducing unnecessary chemical inputs are
plausible conservation measures, but their effects should be tested directly in
replicated studies.
In addition to the absence of spatial replication,
acoustic monitoring may underestimate taxa with low-intensity calls and may not
permit definitive species-level identification of acoustically similar taxa.
The walked-transect design may also generate taxon-specific differences in
detectability. Future studies should include multiple orchard–vineyard pairs
and integrate direct measurements of insect abundance, pesticide exposure,
vegetation structure, landscape composition, and roost availability.
This preliminary case study revealed consistently higher
bat activity, monthly richness, and Shannon diversity at the sampled low-input
citrus orchard than at the sampled intensively managed vineyard. The orchard
also supported a more compositionally diverse assemblage, whereas activity at
the vineyard was dominated by Pipistrellus kuhlii.
These findings indicate that the two agricultural sites differed substantially
in their use by bats during the monitoring period.
Because only one site represented each agricultural
context, these differences cannot be attributed specifically to crop type or
management intensity. They may also reflect site-specific variation in
surrounding landscape, roost proximity, prey availability, microclimate, or
detection conditions. The results should therefore be considered baseline
evidence and a source of hypotheses rather than a general comparison of
orchards and vineyards.
Replicated studies across multiple farms should combine
acoustic monitoring with measurements of insect abundance, pesticide exposure,
vegetation structure, landscape composition, and roost availability. Such work
is needed to determine whether low-input perennial crops consistently
contribute to the conservation of protected bat assemblages in northern African
agricultural landscapes.
Table 1. Characteristics of the selected citrus
orchard and vineyard in the Sebaou Valley, near Draâ Ben Khedda, Tizi Ouzou Province,
northern Algeria.
|
Characteristics |
Site A:
citrus orchard |
Site B:
vineyard |
|
Geographical coordinates |
36.740°
N, 3.958° E |
36.741° N, 3.960° E |
|
Variety |
Orange (Washington Navel) |
Table grape (Red Globe) |
|
Rootstock |
Citrus volkameriana |
41B |
|
Management intensity |
Low-input management |
High-input management |
|
Weeding method |
No weeding |
Chemical weeding |
|
Phytosanitary pressure |
No
pesticide application during the monitoring period; historically low input
use. |
Regular treatments |
|
Type of irrigation |
Drip irrigation |
Drip irrigation |
|
Soil texture |
Sandy clay loam |
Sandy clay loam |
|
Age (years) |
5 |
5 |
|
Area (ha) |
1.0 |
1.5 |
|
Plant spacing (m) |
3 × 4 |
3 × 3 |
Table 2. Acoustic parameters of
bat taxa recorded at the two agricultural study sites in northern Algeria.
Values are presented as mean ± SD.
|
Taxon |
N |
Pulse structure |
SF (kHz) |
EF (kHz) |
FME (kHz) |
D (ms) |
Identification source |
|
Rhinolophus hipposideros |
2 |
FM/CF/FM |
100.06 ± 1.55 |
97.73 ± 0.89 |
115.8 ± 0.62 |
45.3 ± 2.70 |
Ahmim et al. (2020) |
|
Eptesicus isabellinus |
153 |
FM/QCF |
39.58 ± 9.27 |
23.15 ± 1.57 |
27.24 ± 1.28 |
9.54 ± 2.01 |
Ahmim et al. (2020) |
|
Hypsugo savii |
20 |
FM/QCF |
38.48 ± 4.02 |
31.63 ± 2.20 |
33.66 ± 1.17 |
9.01 ± 2.84 |
Disca et al. (2014) |
|
Pipistrellus kuhlii |
222 |
FM/QCF |
63.94 ± 8.16 |
38.76 ± 1.23 |
41.24 ± 1.14 |
5.19 ± 1.50 |
Dalhoumi et al. (2016) |
|
Pipistrellus pipistrellus |
18 |
FM/QCF |
62.9 ± 8.04 |
46.45 ± 3.1 |
50.02 ± 2.37 |
6.58 ± 2.22 |
Russo & Jones (2003) |
|
Plecotus cf. gaisleri |
8 |
FM |
50.24 ± 5.20 |
26.86 ± 1.07 |
33.28 ± 0.94 |
3.11 ± 0.57 |
Dalhoumi et al. (2016); |
N—number of sequences measured in
this study | FM—frequency-modulated | CF—constant-frequency |
QCF—quasi-constant-frequency | SF—start frequency | EF—end frequency |
FME—frequency of maximum energy | D—duration | ms—milliseconds.
Table 3. Results of the negative
binomial regression model (glmmTMB) for bat activity
(total acoustic contacts per pooled one-hour site × month session). Regression
coefficients (β), standard errors (SE), incidence rate ratios (IRR), 95%
confidence intervals (95% CI), and p-values are presented.
|
Predictor |
β |
SE |
z |
p |
IRR (95% CI) |
|
Site: Vineyard vs
Orchard |
-1.538 |
0.256 |
-5.994 |
<0.001 |
0.215 (0.130–0.355) |
|
Month: April vs
March |
-0.742 |
0.489 |
-1.517 |
0.129 |
0.476 (0.182–1.242) |
|
Month: May vs March |
-0.550 |
0.489 |
-1.125 |
0.261 |
0.577 (0.221–1.504) |
|
Month: June vs
March |
-2.671 |
0.656 |
-4.071 |
<0.001 |
0.069 (0.019–0.250) |
|
Month: July vs
March |
-0.392 |
0.492 |
-0.796 |
0.426 |
0.676 (0.257–1.774) |
|
Month: August vs
March |
-0.219 |
0.475 |
-0.461 |
0.645 |
0.803 (0.317–2.039) |
|
Month: September vs
March |
-0.254 |
0.484 |
-0.524 |
0.600 |
0.776 (0.301–2.003) |
|
Month: October vs
March |
0.174 |
0.482 |
0.360 |
0.719 |
1.19 (0.462–3.061) |
|
Month: November vs
March |
-1.287 |
0.515 |
-2.500 |
0.012 |
0.276 (0.101–0.757) |
Table 4. Comparison of monthly taxon richness and
Shannon diversity between the citrus orchard and vineyard. Values represent
median monthly observations. Differences between habitats were assessed using
the paired Wilcoxon signed-rank test.
|
Metric |
Citrus orchard (median) |
Vineyard (median) |
Median of paired monthly
differences |
Statistics (V; p) |
|
Shannon diversity |
0.971 |
0 |
0.656 |
44; 0.0129 |
|
Monthly richness |
3 |
1 |
2 |
28; 0.0213 |
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Table S1. Conservation status and legal protection of
the bat taxa recorded at the two agricultural study sites in northern Algeria.
|
Taxon (authority) — Family |
Common name |
IUCN Red List category |
Global population trend |
CMS listing¹ |
Bern Convention² |
Legal status in Algeria³ |
Conservation relevance |
|
Rhinolophus hipposideros (Borkhausen,
1797) — Rhinolophidae |
Lesser Horseshoe Bat |
Least Concern |
Decreasing |
European populations |
Appendix II |
Protected |
Species with a decreasing global population trend;
acoustically detected only at the orchard site |
|
Eptesicus isabellinus (Temminck,
1840) — Vespertilionidae |
Isabelline Serotine |
Least Concern |
Unknown |
European populations |
Appendix II |
Protected |
Western Mediterranean and North African taxon; one
of the two dominant taxa at the orchard site |
|
Hypsugo savii (Bonaparte, 1837) — Vespertilionidae |
Savi’s Pipistrelle |
Least Concern |
Stable |
European populations |
Appendix II |
Protected |
Widespread Mediterranean species; recorded only at
the orchard site |
|
Pipistrellus kuhlii (Kuhl, 1817) — Vespertilionidae |
Kuhl’s Pipistrelle |
Least Concern |
Increasing |
European populations |
Appendix II |
Protected |
Widespread generalist and the dominant taxon at the
vineyard site |
|
Pipistrellus pipistrellus (Schreber,
1774) — Vespertilionidae |
Common Pipistrelle |
Least Concern |
Stable |
European populations |
Appendix III |
Protected |
Widespread species recorded at both study sites |
|
Plecotus cf. gaisleri Benda
et al., 2004 — Vespertilionidae⁴ |
Long-eared Bat provisionally assigned to Gaisler’s Long-eared Bat |
Not Evaluated for P. gaisleri |
Unknown |
Not confirmed for the North African taxon |
Appendix II |
Protected |
Provisional identification of a probable northern
African endemic taxon; acoustic data alone do not provide definitive species
confirmation |
¹
CMS: Listings follow Appendices I and II of the Convention on the
Conservation of Migratory Species of Wild Animals (CMS 2024). For the families
concerned, Appendix II coverage is generally expressed in relation to European
populations and should not automatically be interpreted as applying to the
Algerian populations recorded in this study.
²
Bern Convention: Listings follow Appendices II and III of the Convention on
the Conservation of European Wildlife and Natural Habitats (Council of Europe
2024). Most microchiropteran bats are included in
Appendix II, whereas Pipistrellus pipistrellus
is included in Appendix III. These listings describe the status of the taxa
under the Convention and should not be interpreted as the source of their legal
protection in Algeria.
³
Algerian protection: National legal status follows Executive Decree No.
12-235 of 24 May 2012 establishing the list of protected non-domestic animal
species (Décret exécutif n° 12-235 du 3 Rajab 1433 correspondant au 24 mai 2012 fixant la liste des espèces animales non domestiques protégées; People’s Democratic
Republic of Algeria 2012).
⁴ Calls assigned to Plecotus
could not be unequivocally identified to species level. Because P. gaisleri is the expected regional
taxon, the records are reported as Plecotus cf. gaisleri. This designation indicates
probable, but not definitive, species identity.
CMS—Convention on the Conservation of
Migratory Species of Wild Animals.