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Article

Effects of Climate on Scorpion Diversity in Arid Ecosystems of the Sahara Desert of Algeria

by
Salah Eddine Sadine
1,
Zineb Souilem
2,
Yacine Belgaid
3,
Abdelwahab Chedad
4,5,
Zineb Djelloud-Souilem
4,5,
Bahri Chebihi
5,
Abdelheq Zouaimia
3,
Zinette Bensakhri
3,
Moussa Houhamdi
3 and
Rabah Zebsa
3,*
1
Laboratoire Valorisation et Conservation des Ecosystèmes Arides (LVCEA), Faculté des Sciences de la Nature et de la Vie et Sciences de la Terre, Université de Ghardaïa, Ghardaïa 47000, Algeria
2
Laboratoire de Mathématiques et Sciences Appliquées (LMSA), Faculté des Sciences et de la Technologie, Université de Ghardaïa, Ghardaïa 47000, Algeria
3
Laboratoire Biologie, Eau et Environnement (LBEE), Faculté SNV-STU, Université 8 Mai 1945 Guelma, 401, Guelma 24000, Algeria
4
Laboratory of Saharan Bio-Ressources: Preservation and Valorisation (BRS), University Kasdi-Merbah, 511, Ouargla 30000, Algeria
5
Directorate of Forest Conservation of Ghardaïa, Ghardaïa 47000, Algeria
*
Author to whom correspondence should be addressed.
Diversity 2023, 15(4), 541; https://doi.org/10.3390/d15040541
Submission received: 26 January 2023 / Revised: 27 March 2023 / Accepted: 3 April 2023 / Published: 9 April 2023
(This article belongs to the Special Issue Diversity of Terrestrial Invertebrate Communities)
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Abstract

:
In desert ecosystems, arthropods such as scorpions are understudied, and sufficient information is still lacking regarding their biodiversity. Specimen collection was carried out over 24 months (2016–2017). This study assessed the phenology, abundance, richness and diversity of scorpion species in arid ecosystems of the Sahara desert of Algeria (Ghardaïa). It examined the potential influence of climate parameters (precipitation, temperature and wind) on activity density, diversity and the phenological distribution of the species among seasons. We identified eight Buthidae species: Androctonus aeneas, Androctonus amoreuxi, Androctonus australis, Buthacus samiae, Buthacus spinatus, Buthacus elmenia, Buthus saharicus and Lissothus chaambi. Androctonus amoreuxi and Androctonus australis were the most abundant and omnipresent species, comprising 54.41% and 33.82% of all species found, respectively. Shannon’s index and the evenness index demonstrated a very poor diversity of scorpions in this region and a poor balance between the number of sampled species. Seasonal variation and climate parameters, i.e., temperature and wind, influenced the number, distribution, and the diversity of scorpions. The number of species found in Ghardaïa Province represent more than 20% of the scorpion species reported in Algeria.

1. Introduction

Biodiversity encompasses all aspects of biological variation on a multidimensional level [1]. It refers to the variety of living organisms, from genes and species to habitats and ecosystems, including the critical roles they perform, that make up the intricate web of life. Anthropogenic pressures such as land-use intensification have precipitated a worldwide biodiversity loss [2], particularly among forest and grassland arthropod species [3,4,5]. In fact, due to their low productivity and diversity [6,7], drylands and deserts have received less scientific attention, despite their critical importance in harboring sensitive endemics and the most endangered species [8,9]. Desert ecosystems are routinely under-documented, and sufficient information and knowledge are still lacking regarding their biodiversity [10,11].
Scorpions are one of the most important taxa for ecological, conservation and biogeographic studies [12,13]. They are among the oldest known Arthropods (more than 450 million years) [14,15]; they have lived through all geological eras and are particularly abundant in deserts and unpopulated areas due to their adaptability and their ecological plasticity [16,17].
To date, zoologists have described more than 2740 species [18], of which more than 50% have been documented from the Neotropical region [19] and over 1000 have been described during the past two decades [20]. Scorpions have radiated into all nonboreal habitats, including deserts, savannas, grasslands, temperate forests, tropical forests, rain forests, the intertidal zone, and snow-covered mountains over 5500 m in altitude [21].
Many environmental factors can influence the diversity and abundance of scorpions in most ecosystems, such as the soil type, topography, hydrology, food resources, and especially, temperature and precipitation [13,22,23,24,25,26,27,28,29,30], which are the important determinants of the general geographical range of cryptozoic species [12,29,31,32,33,34].
Scorpions are primarily solitary and sedentary arthropods and live preferentially in microhabitats [35]. The processes of habitat selection usually involve responses to environmental conditions that promote the growth, survival and reproductive success of this species [36,37,38,39].
Scorpion fauna of Algeria are truly native, and the endemic species represent more than 59% [40,41]. More than 49 species can be found [42,43], and the species are assembled in three major groups: scorpions of Algerian–Tunisian compartment (Mediterranean area), those of Saharan sector and scorpions of Saharo-mountainous sector [44]. Nevertheless, scorpion diversity is particularly high in deserts and arid habitats [45,46] because the temperature has an effect on scorpion communities [22,47,48,49], profoundly modulating several physiological processes such as digestion, oxygen consumption [50,51], growth [52] and movement performance in ectothermic animals such as scorpions [53].
In this context, we performed an ecological study on scorpions in central Algeria. This research used Ghardaïa Province as a study case with the following objectives: (i) to establish a faunistic list of scorpion species; (ii) to determine the phenological pattern of species; (iii) to study the effect of different climate factors in arid ecosystems (temperature, precipitation and wind) on scorpion diversity, distribution and phenology.

2. Materials and Methods

2.1. Study Site

Our study was conducted in the north of the central Algerian Saharan region, specifically in Ghardaïa Province (29°19′ N−32°57′ N, 02°03′ E−04°54′ E) (Figure 1a), which has an area of ca. 86,560 km2. The average altitude of the main reliefs is 520 m. Geomorphological features constitute wadis and reg formations (Figure 1b,c) [54]. The region is characterized by a dry Saharan climate, with extreme thermal amplitudes between the day and the night that reach 15–16 degrees [55]. The coldest month is January, with a minimum temperature of 6.2 °C, whereas the hottest month is July, with a maximum temperature of 41.8 °C.
Rainfall is extremely low in the region of Ghardaïa, with an average of 80.2 mm per year. Air humidity is rather low, with a maximum humidity of 55.5% in December and a minimum of 21.6% in July [56]. An analysis of dry periods over several years attested that 11 months are dry, ranging from February to December; only a short and slightly more humid period can be experienced in January.

2.2. Scorpion Sampling

As scorpions are nocturnal predators, preliminary visits to the sites during the nighttime were carried out to confirm the existence of scorpions. However, for practical and security reasons, the collection of scorpions was carried out during the daytime period. Samples were taken monthly over 24 months, between January 2016 and December 2017, and in the same locations. We carried out 24 field trips (each lasting 4 h) during the four seasons, and our capture followed a strategy of systematically searching the shelters, which were chosen at random or were suspected of harboring the scorpions during the visual search in each field trip. All samples were separately preserved in 70° ethanol.
The collected specimens were taken to the Laboratory of Zoology, University of Ghardaïa. They were then identified to their genus and species based on morphological characters, using a binocular magnifying glass. The nomenclature and taxonomy of species were made based on up-to-date references, following measurements [57], Trichobothrial notations [58] and morphological terminology [44,59].

2.3. Data Analysis

Statistical analyses were carried out with R 4.0.2 [60]. Ecological indices were estimated based on the scorpion fauna data to characterize biodiversity in the different sampling season during the study period, which was evaluated by calculating: (i) abundance (Ni), the total number of caught individuals of each species; (ii) the relative abundance (RA%) for each species, which was determined as the ratio of the number of individuals for each species divided over the total number of individuals recorded; (iii) the occurrence frequency©, which was calculated as follows: C = (p × 100)/N, where C represents the constancy % for each species, p represents the number of samples in which the species is present, and N represents the total number of samples [61]. According to their frequency of occurrence, four species groups are distinguished by Bigot and Bodot [62]: very accidental species (Vac), represented by an occurrence of less than 10%; accidental species (Acc), whose occurrence varies between 10 and 24%; common species (Cmt), which are present in 25–49%; constant species (Cst), which are present in 50–57% and omnipresent species (On), whose occurrence is more than 75%. The biodiversity of scorpions was also assessed by (iv) species richness (S), which represents the total number of species identified. Additionally, (v) Shannon’s index (H′ = −∑(pi × log2pi)), the maximum of Shannon index: Hmax = log2(S); Ratio N/S; Ratio SRI/S; Simpson reciprocal index: SRI = (1/D), with D = Σ(Ni(Ni – 1)/N(N – 1)), where (Ni) represents the number of individuals of a given species of rank (i); and (vi) evenness (evenness = H′/log2Sobs) were applied for measuring scorpion diversity in each sampled season period, based on the relative density pi of the ith species [61].
The analysis of the similarity between species of scorpions and seasons was carried out by calculating several similarity indices with the software EstimateS [63]. These included qualitative indices (Jaccard and Sørensen) and quantitative indices: the Morisita–Horn index and Bray–Curtis index. Moreover, correlation matrices and a Venn diagram were drawn up in order to illustrate all possible relationships between diversity parameters of scorpions and different scorpion species according to different seasons in the Ghardaïa region.
Generalized linear mixed effect models with Poisson error distribution were applied to assess for the effect of climate variables (T, PP and W) and RA as explanatory variables for the abundance of all species separately, with date and sites taken as random effects (to account for overdispersion). Another generalized linear mixed effect model was conducted to analyze the variation in diversity parameters across the season. The model included the season (cold, pre-hot, hot and post-hot) as explanatory variable, and the number of individuals and diversity index (SR, N:S, Shannon’s index, etc.) were included as response variables. All models were fitted using the R package, lme4 [64]. Values presented hereafter are shown as the mean ± SD.

3. Results

In our survey, a total of 408 scorpions were collected in the Ghardaïa region, during the period from 2016–2017, with 229 and 179 individuals, respectively.

3.1. Taxonomic Composition, Relative Abundance and Occurrence

Based on the identification keys mentioned previously, our study revealed that the scorpion assemblage in Ghardaïa region is composed of eight species. All species belong to four genera in the Buthidae family (Table 1).
The genus Androctonus was represented by three species: A. aeneas, A. amoreuxi and A. australis. The Buthacus genus was also presented by three species: B. elmenia, B. samiae and B. spinatus. The genera Buthus and Lissothus were presented each by a single species, B. saharicus and L. chaambi, respectively. Among the four genera found in the Ghardaïa region, the Androctonus species were the most abundant (RA = 91.18%), while the other genera were represented with a very low abundance (less than 4.4%). Androctonus amoreuxi and A. australis were the most abundant species with (54.41 and 33.82%), respectively, and they were classified as omnipresent species. The other species are represented with a very low abundance with different occurrences: Lissothus chaambi was ranked as a constant species, while Androctonus aeneas, B. samiae and B. saharicus were classified common species. Two species, B. elmenia and B. spinatus, were classified as very accidental species in our study.

3.2. Species Phenology

The phenogram (Figure 2) shows that the monthly temporal abundance of the scorpion species recorded in Ghardaïa region was very variable during the different months in 2016 and 2017.
In our study, most of the species were present in the pre-hot (21.32%) and hot seasons (59.31%). Androctonus amoreuxi and A. australis were the dominant species over the 24 months (2016–2017). Lissothus chaambi and B. saharicus were present throughout the year but demonstrated a low abundance. Similarly, A. aeneas was present throughout the year but was absent the in the cold season. Contrariwise, B. samiae was absent in the post-hot season. However, B. elmenia and B. spinatus were found only in February and March (the pre-hot period) of 2017.
The results of the linear mixed models (GLMM) applied to reveal the effect of the ecological descriptors parameters (temperature, precipitation and wind) on the abundance of each scorpion species separately are represented in Table 2. There was a significant positive relationship between wind and the abundance of the three species A. amoreuxi, A. australis and B. elmenia (p < 0.05). Only A. amoreuxi showed a potential density-dependent population, which was revealed by the negative relationship between the number of individuals and the relative abundance of the species (p < 0.05). There was a significant positive effect of temperature on the abundance of A. australis (p < 0.05), but with a significant negative relationship with B. samiae (p < 0.05). Only the number of individuals of B. saharicus increased significantly with the increase in precipitation (p < 0.05). However, the abundance of A. aeneas, B. spinatus and L. chaambi was independent of these climatic factors (p > 0.05).

3.3. Variation of Diversity Parameters

3.3.1. Number of Individuals

The number of individuals captured varied depending on the seasons. In fact, the highest number of individuals captured was recorded in the pre-hot and hot seasons, with an average of 21.75± 9.11 and 20.17 ± 8.73 individuals, respectively. The lowest number was recorded in winter, with an average of 8.75 ± 1.26 individuals (Figure 3).

3.3.2. The Ecological Indices according to Seasons

During the different months (2016–2017), we identified a total of eight species. Figure 3 shows that the specific richness varied according to the seasons. The pre-hot seasons contained five species, while other seasons contained three. We note that the specific richness showed averages ranging from 5.25 ± 0.50, recorded in the pre-hot seasons, to 3.00 ± 1.15, recorded in the post-hot season. The other two seasons (cold and hot) recorded 3.75 ± 0.96 and 3.33 ± 0.78 species, respectively.
During the study period, the most important mean values of the ratio N:S were recorded in the hot seasons (6.29 ± 2.87). The post-hot and pre-hot seasons are represented by 4.31 ± 2.25 and 4.07 ± 1.34, respectively. The lowest value of the ratio N:S was recorded in the cold seasons, with an average of 2.43 ± 0.65 (Figure 3).
In cold seasons, the Shannon diversity index (H′) was mostly recorded with a mean value of 1.62 ± 0.32 bits. The values of this index in the pre-hot, hot and post-hot seasons were 1.53 ± 0.35, 1.31 ± 0.25 and 1.18 ± 0.40, respectively.
The most important value, the average of Shannon’s maximum index, was recorded in the pre-hot (2.39 ± 0.13) season, followed by the cold seasons (1.87 ± 0.36) and hot seasons (1.70 ± 0.34). The post-hot seasons are represented by a mean value of 1.50 ± 0.58 (Figure 3).
Evenness recorded an overall average value of 0.77 ± 0.11. The most important evenness value was recorded in the cold (0.86 ± 0.02) and post-hot seasons (0.80 ± 0.09). The hot seasons are represented with an average of 0.78 ± 0.09, while the pre-hot seasons recorded the lowest evenness, with an average of 0.64 ± 0.16 (Figure 3).
The general mean value of SRI during the study period was 2.27 ± 0.47. According to Figure 3, the most important average values were recorded in the cold seasons (2.69 ± 0.58), followed by the pre-hot seasons (2.27 ± 0.63) and the hot seasons (2.23 ± 0.34). The post-hot seasons were represented by the lowest average value of 1.97 ± 0.45 (Figure 3).
The overall mean value of the ratio (SRI:S) recorded a value of 0.65 ± 0.15. We note that the cold and post-hot seasons recorded were almost equal, with 0.72 ± 0.06 and 0.70 ± 0.17, respectively. The value in the hot seasons was 0.69 ± 0.12. The lowest mean value of the ratio was recorded in the pre-hot seasons (0.44 ± 0.14) (Figure 3).

3.4. Relationships between Diversity Parameters

The relationships between the diversity indices are summarize in Figure 4, which shows that there is generally a significant positive correlation (Pearson correlation: p < 0.05) between (S) and the other descriptors except (N), and there was a negative relationship between (N:S). A significant positive relationship was detected between N and N:S but was negative with E.

3.5. Effects of Climate on Variations of Biodiversity Parameters

The linear mixed models (GLMM) applied for the effects of climate on variations of the biodiversity parameters in the scorpions of the Algerian Sahara are represented in Table 3.
Our GLMM models showed that there was a highly significant effect of temperature on diversity indices H and Hmax in which the values decreased with the increase in the latter (Table 3). A significant positive relationship was detected between the number of scorpion individuals and the speed of the wind. Moreover, the ratios N:S and SRI:S varied significantly with temperature, and they increased with the increase in temperature. For the rest of the parameters, no effect was revealed (p > 0.05) (Table 3).

3.6. Distribution and Similarity Analysis of Scorpions According to the Seasons

According to the Venn diagram (Figure 5), four species were common between seasons, i.e., A. amoreuxi, A. australis, B. saharicus and L. chaambi. The B. elmenia and B. spinatus were characteristic of the pre-hot season. B. samiae was found in all three seasons (cold, pre-hot and hot seasons), while A. aeneas was common between the three hot seasons (pre-hot, hot and post-hot seasons).
The similarity calculation between the different seasons (Table 4) showed that the most important similarity was found between the cold and hot seasons, with a 83.3 and 90.9% similarity for the Jaccard and Sorensen, respectively, whereas the Morisita–Horn index showed a 99.4% similarity between the hot and post-hot seasons. The Bray–Curtis dissimilarity index showed that the greatest dissimilarity existed between the cold and post-hot (70.1%) seasons, while the least significant dissimilarity exists between the cold and hot (25.3%) seasons (Table 4).

4. Discussion

Our survey is the first of its kind on scorpions in Algeria’s arid ecosystem (Ghardaïa), which is characterized by an arid climate with hot summers. During the study period, we documented eight scorpion species, accounting for more than half of the scorpions in the northern Algerian Sahara [65], with good congeneric coexistence [66,67,68,69,70].
Most of those species are recorded as Saharan endemic scorpions [40,44,71]. In neighboring regions of Ghardaïa, [72] identified eight scorpions species in El-Oued and nine scorpion species in Ouargla, [40]. In the Algerian forest and mountainous ecosystems, up to ten scorpion species were identified in Sidi Bel Abbes, northwest Algeria [73] and eight scorpion species were identified in the Ouarsenis massif of Tissemsilt, northwest Algeria [23]. However, it seems that the species richness of scorpions was lower in other regions, such as the Khenchela region [74] and Tebessa region [42], with only five species identified. During the study period, five new records of species were discovered for the region, i.e., B. elmenia, B. samiae, B. spinatus, B. saharicus and L. chaambi, reflecting an important endemism for this region. Lourenço et al. [75] suggested in particular the presence of micro endemic populations of scorpions in Ghardaïa deserts.
Among the eight species found in the Ghardaïa region, A. amoreuxi is the most abundant species, followed by A. australis. Similar results were indicated in Sadine [40], with a rate of 42% for these two species. However, many other authors mentioned that A. australis is the most abundant and the most widespread species in the Algerian Sahara [40,43,71,76,77]. As Androctonus australis is limited to North Africa, in particular to Algeria, Egypt, Libya, Sudan and Tunisia [78], this species remained the most abundant (RA = 31.25%) in the Misurata region, North Libya [79]. This species is also widespread in different eco-geographical regions in Egypt [80]. The other species, i.e., A. aeneas, B. elmenia, B. samiae, B. saharicus, B. spinatus and L. chaambi, ranked as rare species and are represented by ratios of less than 0.5%. The same results were reported in previous studies with less than 3% [40,76,77].
The relative abundance of Ghardaïa scorpions is variable in months and shows an annual activity with a potential dynamic observed in some months (the hot period). However, they have a slight presence in the cold period. According to Sadine [81], scorpions in Ouargla (Algerian Sahara) are active in the spring and summer, and they begin their winter diapause (hibernation under shelters) at the beginning of autumn. While some species may retain their ability to be active during the cold season, this only justifies a minor presence during the cold season [82].
Nonetheless, some scorpions undergo semi-hibernation; if disturbed, they maintain the whole of their resources, which they demonstrate by being alert [83,84].
Furthermore, the abundance of certain species, such as A. australis and A. amoreuxi, can be deduced from average number of offspring for these scorpions; for example, A. australis can produce more than 130 offspring [71]. While, in Morocco, Lissothus occidentalis produces 2 to 14 pullus (offspring) [85].
Both A. amoreuxi and A. australis were classified as omnipresent species during the study period. The former was also reported as omnipresent in Morocco [86] and Egypt [80,87], but it was ranked as very accidental species (Occ = 8.33%) in the North of Libya [88]. The second species was as omnipresent in all previous countries. The constant species category is represented by L. chaambi. A. aeneas, B. samiae and B. saharicus, which were classified as common species. Two species, B. elmenia and B. spinatus, were classified as very accidental species in our study.
Only three species (A. amoreuxi, A. australis and L. chaambi) showed annual activity in our study area, with a potential dynamic between May and August and a slight absence during the cold periods. This significant increase in population can be attributed to an increase in the number of males preparing for mating [40,84]. Buthidae, like southern hemisphere scorpions, mate in the autumn and begin their gestation period from November to April, lasting 5.2 ± 2.3 months [13,89]. The mating behavior of A. australis in the Ouargla region begins in late August and lasts until mid-October, when the climate conditions are characterized by relatively low nighttime temperatures, ranging from 18 to 20 °C, and frequent winds with high humidity [84]. Sadine et al [40] reported that the phenological cycle of A. amoreuxi is nearly identical to that of A. australis, with a minor difference in the favorable period for mating (coupling).
Many environmental factors may influence scorpion activity in arid ecosystems [26]. In ectothermic animals such as scorpions, temperature profoundly modulates several physiological processes including, but not limited to, digestion, oxygen consumption [50,51], growth [52] and movement performance [53]. The temporal distribution of the species reveals that the pre-hot and hot seasons are the best time to collect scorpions. This can be explained by the fact that scorpion activity begins during this season, which corresponds to scorpion reproduction and an increase in their locomotor activity [90]. The number of individuals harvested in winter was very low (an average of 8.75 ± 1.26 individuals), which is explained by the fact that this season corresponds to the scorpions’ lowest activity [37].
Hotspots of scorpion species richness at continental or regional scales are associated with areas of climatic, topographic and geological complexity [29]. Climate factors, substrate (soil hardness and texture; amount of stone or litter cover), and vegetation physiognomy all influence scorpion assemblages and their feeding resources at the local scale [29,91,92]. Indeed, Haghani et al. [93] discovered that ecological factors have a significant impact on the families Buthidae and Scorpionidae.
The results obtained regarding species diversity show that climate variables have an effect on scorpion biodiversity. Previous research indicated that temperature and precipitation play an important role in the distribution and beta diversity of scorpion species assemblages [13,25,29,92,94,95]. Algeria, for example, with its vast geographical extent and diverse ecosystems, attests to a high level of diversity, with more than 49 species of scorpions [42,43].
This study allowed us to explore this region in order to better direct our efforts in the management and conservation of scorpion populations. We hope that this pioneer study will inspire scientists in North Africa to use this fascinating biological system in future research because this taxon, with its relatively limited dispersion and long life cycles, is an excellent tool for studying patterns and processes related to biological gradient diversity [29,92,94,96,97,98].

5. Conclusions

This study focused on scorpion diversity in arid ecosystems of Algeria’s Sahara Desert. Despite the conditions of this region, we documented eight species during our survey over the two years of the study (2016–2017). Our findings demonstrate a modest level of diversity of scorpions in the Algerian Sahara compared to other regions in the world [99,100]. The study revealed temporal variations in scorpion biodiversity and demonstrated that several parameters influence the distribution of the scorpion fauna. Indeed, seasonal changes may cause variations in feeding resources which affect the temporal distribution and abundance of scorpion species. Orthochirus innesi, the ninth scorpion species, was recently recorded in central Algeria [101], with a new locality in Ghardaïa palm groves. It would be preferable to increase the number of studies that could lead to new discoveries regarding scorpion biodiversity in this understudied area.

Author Contributions

Conceptualization, S.E.S., Z.S., Y.B., A.C., Z.D.-S. and B.C.; methodology, S.E.S., Z.B. and M.H.; software, R.Z.; validation, S.E.S., M.H. and A.Z.; formal analysis, R.Z.; data curation, Z.B. and A.Z.; writing original draft preparation and writing—review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

This work is part of Project PNR-2021, on the scorpion diversity of central Algeria, sponsored by The Thematic Agency for Research in Health and Life Sciences (ATRSSV), Algeria.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The datasets used and analyzed during the current study are available from the corresponding author upon reasonable request.

Acknowledgments

We gratefully acknowledge H. Chenchouni for his help with the statistical analysis. We are thankful to the reviewers for their constructive comments, which improved an earlier version of the manuscript. We are grateful to all students and colleagues who contributed to the fieldwork. This research was supported by MESRS Algeria.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. Study area. (a) Map of Algeria showing the Ghardaïa region; (b) reg formations and “desert pavement”; (c) wadi bed.
Figure 1. Study area. (a) Map of Algeria showing the Ghardaïa region; (b) reg formations and “desert pavement”; (c) wadi bed.
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Figure 2. Phenogram representing monthly abundance of scorpions in the Ghardaïa region during the two years of study (red = 2016; green = 2017).
Figure 2. Phenogram representing monthly abundance of scorpions in the Ghardaïa region during the two years of study (red = 2016; green = 2017).
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Figure 3. Box plots representing the different ecological indices applied to scorpion assemblages sampled during different seasons.
Figure 3. Box plots representing the different ecological indices applied to scorpion assemblages sampled during different seasons.
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Figure 4. Correlation matrix between diversity indices of scorpions in the Sahara Desert of Algeria (Values above diagonal are Pearson’s correlation coefficients, and figures below diagonal represent p-values).
Figure 4. Correlation matrix between diversity indices of scorpions in the Sahara Desert of Algeria (Values above diagonal are Pearson’s correlation coefficients, and figures below diagonal represent p-values).
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Figure 5. Venn diagram explaining the distribution of number of scorpion species according to seasons in the Ghardaïa region, Sahara Desert of Algeria.
Figure 5. Venn diagram explaining the distribution of number of scorpion species according to seasons in the Ghardaïa region, Sahara Desert of Algeria.
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Table
Table 1. Species list, abundance and occurrence frequency of scorpion species captured in the Ghardaïa region, Algeria (2016–2017).
Table 1. Species list, abundance and occurrence frequency of scorpion species captured in the Ghardaïa region, Algeria (2016–2017).
Genus (RA in %)SpeciesTraitsSeasonsOverall
ColdPre-HotHotPost-Hot
Androctonus (91.18)A. aeneas C.L. Koch, 1839Ni—56112
RA [%]—1.231.470.252.94
Occ—7541.72537.5
Scale—OmnComComCom
A. amoreuxi (Audouin, 1826)Ni165712524222
RA [%]3.9213.9730.645.8854.41
Occ100100100100100
ScaleOmnOmnOmnOmnOmn
A. australis (Linnaeus, 1758)Ni10159716138
RA [%]2.453.6823.773.9233.82
Occ100100100100100
ScaleOmnOmnOmnOmnOmn
Buthacus (2.45)B. elmenia Lourenço and Sadine, 2017Ni—2——2
RA [%]—0.49——0.49
Occ—50——8.3
Scale—Cst——Vac
B. samiae Lourenço and Sadine, 2015Ni242—8
RA [%]0.490.980.49—1.96
Occ507516.7—29.2
ScaleCstOmnAcc—Com
B. spinatus Lourenço, Bissati and Sadine, 2016Ni—2——2
RA [%]—0.49——0.49
Occ—50——8.3
Scale—Cst——Vac
Buthus (1.47)B. saharicus Sadine, Bissati and Lourenço, 2016Ni21216
RA [%]0.490.250.490.251.47
Occ502516.72525
ScaleCstComAccComCom
Lissothus (4.41)L. chaambi Lourenço and Sadine, 2014Ni5110218
RA [%]1.230.252.450.494.41
Occ752558.35054.2
ScaleCstComCstCstCst
Total Ni358724244408
RA [%]8.5821.3259.3110.78100
RA, relative abundance (%); Ni—total number of caught individuals; Occ—occurrence frequency; Acc—very accidental species; Com—common species; Cst—constant species; Omn—omnipresent species.
Table
Table 2. GLMM testing effects of climate on variations of scorpion species abundances in the Sahara Desert of Algeria.
Table 2. GLMM testing effects of climate on variations of scorpion species abundances in the Sahara Desert of Algeria.
VariablesEstimateS.E.z-ValuepSig.EstimateS.E.z-ValuepSig.
A. amoreuxi A. australis
Intercept1.6050.5233.0680.002**−0.0920.729−0.1270.899ns
T−0.0080.009−0.8430.399ns0.0350.0122.9700.003**
PP−0.0260.023−1.1060.269ns−0.0550.036−1.5260.127ns
W0.0740.0272.6900.007**0.0790.0352.2450.025*
RA−0.1250.054−2.2980.022*−0.0310.072−0.4270.670ns
A. aeneas B. elmenia
Intercept−0.4812.180−0.2210.825ns−10.6915.408−1.9770.048*
T−0.0520.040−1.2940.196ns−0.1430.096−1.4860.137ns
PP−0.1190.165−0.7220.471ns0.0060.2520.0260.980ns
W0.0960.1200.8020.423ns0.7690.3392.2670.023*
RA−0.2040.236−0.8630.388ns0.0470.3180.1470.883ns
B. samiae B. spinatus
Intercept3.7522.6541.4140.158ns−3.1403.332−0.9420.346ns
T−0.1980.094−2.1000.036*−0.1030.068−1.5020.133ns
PP0.0480.1440.3370.736ns−0.2940.371−0.7910.429ns
W−0.0390.150−0.2580.797ns0.2740.1921.4260.154ns
RA−0.4690.284−1.6490.099ns0.0850.2610.3280.743ns
B. saharicus L. chaambi
Intercept−3.2223.643−0.8840.377ns−0.5131.802−0.2850.776ns
T−0.0950.092−1.0400.298ns−0.0330.035−0.9560.339ns
PP0.2400.1211.9930.046*0.0680.0621.1100.267ns
W0.1640.1750.9360.349ns0.0540.0940.5760.565ns
RA0.3250.3400.9560.339ns0.0240.1820.1330.894ns
(T—temperature; PP—precipitation; W—wind; RA—relative abundance (%); **—p < 0.001; *—p ≤ 0.05; ns—p > 0.05).
Table
Table 3. GLMM testing effects of climate on variations of biodiversity parameters in scorpions of the Algerian Sahara.
Table 3. GLMM testing effects of climate on variations of biodiversity parameters in scorpions of the Algerian Sahara.
NEstimateS.E.z-ValuepSig.SEstimateS.E.z-ValuepSig.
Intercept2.1630.3885.579<0.001***Intercept1.7850.7912.2570.024*
T−0.0010.007−0.1760.861nsT−0.0290.015−1.9250.054ns
PP−0.0330.017−1.8610.063nsPP0.0150.0310.4770.633ns
W0.0660.0203.2510.001**W0.0140.0420.3300.742ns
RA−0.1040.040−2.6160.009**RA−0.0460.083−0.5560.578ns
N:SEstimateS.E.t-valuepSig.HEstimateS.E.t-valuepSig.
Intercept−1.0873.418−0.3180.758nsIntercept2.0870.4175.001<0.001***
T0.1490.0572.6000.018*T−0.0250.007−3.5540.002**
PP−0.1310.131−1.0040.345nsPP0.0130.0160.8410.429ns
W0.2220.1721.2930.217nsW−0.0070.021−0.3180.757ns
RA−0.0730.340−0.2140.839nsRA−0.0450.042−1.0680.318ns
HmaxEstimateS.E.t-valuepSig.EEstimateS.E.t-valuepSig.
Intercept2.7340.4945.534<0.001***Intercept0.7630.1804.236<0.001**
T−0.0410.008−5.074<0.001***T0.0040.0031.1500.271ns
PP0.0110.0190.5840.577nsPP0.0000.0070.0560.963ns
W0.0130.0240.5110.618nsW−0.0080.009−0.7910.461ns
RA−0.1000.049−2.0710.059nsRA0.0170.0180.9320.441ns
SRIEstimateS.E.t-valuepSig.SRI:SEstimateS.E.t-valuepSig.
Intercept3.0420.7174.240<0.001***Intercept0.4890.2232.1990.050*
T−0.0260.013−2.0990.052nsT0.0100.0042.6400.017*
PP0.0220.0280.7990.490nsPP−0.0030.009−0.2940.790ns
W−0.0100.038−0.2670.800nsW−0.0070.012−0.6400.540ns
RA−0.0480.073−0.6530.573nsRA0.0220.0230.9920.375ns
T—temperature; PP—precipitation; W—wind; RA—relative abundance (%); Sig.—probability significance; ***—p < 0.001; **—p < 0.01; *—p ≤ 0.05; ns—p > 0.05.
Table
Table 4. Similarity analysis (in %) between number of scorpions and seasons in the region of Ghardaïa, Algeria.
Table 4. Similarity analysis (in %) between number of scorpions and seasons in the region of Ghardaïa, Algeria.
First SampleSecond SampleJaccard ClassicSorensen ClassicMorisita–HornBray–Curtis
ColdPre-hot62.576.990.349.2
ColdHot83.390.995.725.3
ColdPost-hot5066.79470.1
Pre-hotHot7585.791.549.2
Pre-hotPost-hot5066.794.163.6
HotPost-hot66.78099.429.6
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Sadine, S.E.; Souilem, Z.; Belgaid, Y.; Chedad, A.; Djelloud-Souilem, Z.; Chebihi, B.; Zouaimia, A.; Bensakhri, Z.; Houhamdi, M.; Zebsa, R. Effects of Climate on Scorpion Diversity in Arid Ecosystems of the Sahara Desert of Algeria. Diversity 2023, 15, 541. https://doi.org/10.3390/d15040541

AMA Style

Sadine SE, Souilem Z, Belgaid Y, Chedad A, Djelloud-Souilem Z, Chebihi B, Zouaimia A, Bensakhri Z, Houhamdi M, Zebsa R. Effects of Climate on Scorpion Diversity in Arid Ecosystems of the Sahara Desert of Algeria. Diversity. 2023; 15(4):541. https://doi.org/10.3390/d15040541

Chicago/Turabian Style

Sadine, Salah Eddine, Zineb Souilem, Yacine Belgaid, Abdelwahab Chedad, Zineb Djelloud-Souilem, Bahri Chebihi, Abdelheq Zouaimia, Zinette Bensakhri, Moussa Houhamdi, and Rabah Zebsa. 2023. "Effects of Climate on Scorpion Diversity in Arid Ecosystems of the Sahara Desert of Algeria" Diversity 15, no. 4: 541. https://doi.org/10.3390/d15040541

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