Keywords
MaxEnt Model, Niche Assessment, Species Distribution Models
MaxEnt Model, Niche Assessment, Species Distribution Models
Climate change has exerted significant biological, spatial and temporal effects on terrestrial habitats1,2. The increasing CO2 trend during the past decades will continue for several decades, which will likely have major effects on animal and plant species3. Additionally, the inherent fragility of dry lands made them particularly vulnerable to climate change, even a small change in temperature and rainfall patterns may cause serious threat to their biodiversity4. IIED5 suggests that dry-land regions are projected to significant climate changes, but there still exist substantial variability and uncertainty in these estimates based on different scenarios. Many of the important United Arab Emirates (UAE) plant species, for instance, may be facing major challenges for their survival. Many of such native species are becoming restricted in their distributions because of overgrazing and habitat destruction. In addition to the natural prolonged drought, human activities are also threatening many native plants6 and their conservation7. The IPCC report8 also projected that deserts are going to become hotter and drier. Additionally and in relation to the Arab region, climate change-induced desertification has been prolonged, critically affecting the vulnerability of the local population. The projected warming would lead to seasonal changes in precipitation, evapotranspiration and radiation Hulme and Jenkins9. These changes can largely affect the distribution of species and habitat composition10. Thus, the implications of climate change for biodiversity have led to the use of bioclimatic models in order to forecast the species range shift under future climate change scenarios Araújo and New11.
Both habitat quality and diversity have significant impact on species’ distribution and richness within the environment12. Ecological envelope models or species distribution models (SDMs) have been widely used for projecting the habitat shift and distribution induced by climate change13. These models are also used to understand biological processes14 and climate change impacts on biodiversity15–17. Understanding the future distribution of climatically suitable habitat allows the conservation planners to assess the vulnerability of species and ecosystem to climate change18,19. The species occurrence records alone may not be sufficient for building robust conservation strategy in light of climate change. While combining occurrence records with an environmental suitability model can be a promising tool for to assess species distribution and the potential impacts of climate change20,21.
Maximum entropy (MaxEnt) is one of the robust environmental suitability ecological niche/ ecological envelope) models for assessing species habitat suitability changes under the ongoing and future environmental challenges22–24. MaxEnt is a species distribution model (SDM)25 and it is used for the present study. The hypothesis is that shifting, contracting and expanding species habitats are affected by the ongoing climate change. For this, habitat suitability changes of the selected three species (i.e. Calotropis procera, Prosopis cineraria and Ziziphus spina-christi) in the current vs the future climate scenarios was modeled.
This study was conducted in the UAE. It covers a total land area of approximately 82,925 km2, which was used as the basis for calculating the species distribution areas. The study area boundaries was converted to shape files and and then used to extract the environmental and climatic variables.
Calotropis procera, belonging to the Asclepiadaceae family, locally known as ash harbor or milkweed. It is a poisonous shrub that can cause harm to humans if its latex touches the skin or eyes. The latex is a milky white substance present within the shrub, and is released if the shrub was cut26.
Prosopis cineraria is considered the national tree of UAE and belongs to the Fabaceae family. It can withstand a wide range of temperature, heat and drought and also can tolerate saline soil conditions. It commonly exists in India and Pakistan in large numbers. In the UAE it is found on the inland sand plains and low dunes of the Northern Emirates and eastern rim of Abu Dhabi emirate, and appears occasionally in wadi beds of the Hajar mountain range27.
Ziziphus spina-christi, belongs to the Rhamnaceae family, is a species indigenous to the Arabian Peninsula. This species is known for delayed seed germination, which is due to the hard coat of its seeds, therefore lowering its ability to compete with other plants over resources. This tree has different importance to people and animals. The honey produced from its flowers is very valuable and its leaves has various medicinal uses. The leaves also contain various flavonoids, saponin, alkaloids, and allelochemicals.
We have extracted current and future 19 bioclimatic variables from the WorldClim website28 and tested for colinearity using the SDM tool in ArcGis 10.129 (an open-source alternative is QGIS). Additionally, variables included were land use cover and elevation data for the study area. Even though the variables were highly correlating we have selected nine bioclimatic variables and the edaphic variables for the model development. Representative concentration pathway (RCP)4.5 was selected and for 2050s and 2070s, the Community Climate System Model version 4 climate model was used to download the bioclimatic variables. RCP4.5 stabilizes radiative forcing by 210030. It includes global emissions of greenhouse gases and land-use-land-cover. A resolution of approximately 1 km2 (30 arc-seconds) was used for this predictive assessment.
All three species’ presence-only data was used and each species had 100 georeferenced location data. For the selected species, the model was trained using current bioclimatic data and projected using future bioclimatic variables. To visualize the bioclimatic suitability range shift in the future projection, this study has used clamping module while running the model. Final outputs of the model predictions were exported to ArcGIS 10.1 for further analysis. The exported model outputs were converted to raster files and then the predicted habitat suitability ranges were reclassified using three pre-defined probability classes: high, medium and low. Other model settings were kept unmodified and the Jackknife method was used to assess the variable contribution to the habitat distribution. Model fit was assessed using the area under the curve (AUC) values.
In addition to land use cover and elevation data, 19 variables were extracted from the WorldClim database31, which is a set of global climate layers generated through interpolation of climate data from weather stations on a 30 arc-seconds grid (about 1 km2 resolution). The 19 environmental variables are coded as follows:
1. BIO1: Annual Mean Temperature
2. BIO2: Mean Diurnal Range (Mean of monthly (max temp - min temp))
3. BIO3: Isothermality (BIO2/BIO7) (* 100)
4. BIO4: Temperature Seasonality (standard deviation *100)
5. BIO5: Max Temperature of Warmest Month
6. BIO6: Min Temperature of Coldest Month
7. BIO7: Temperature Annual Range (BIO5-BIO6)
8. BIO8: Mean Temperature of Wettest Quarter
9. BIO9: Mean Temperature of Driest Quarter
10. BIO10: Mean Temperature of Warmest Quarter
11. BIO11: Mean Temperature of Coldest Quarter
12. BIO12: Annual Precipitation
13. BIO13: Precipitation of Wettest Month
14. BIO14: Precipitation of Driest Month
15. BIO15: Precipitation Seasonality (Coefficient of Variation)
16. BIO16: Precipitation of Wettest Quarter
17. BIO17: Precipitation of Driest Quarter
18. BIO18: Precipitation of Warmest Quarter
19. BIO19: Precipitation of Coldest Quarter
The MaxEnt model Version 3.4.125 was used for this assessment of the three different tree/shrub species distribution. MaxEnt has been used to model species niches and distributions through machine-learning techniques referred to as maximum entropy modeling17,32. It is very robust in cases of small sample size of species distribution and locations17.
MaxEnt uses a combination of a set of environmental (e.g. climatic) grids and geo-referenced presence points for the species under investigation33. The model expresses a probability distribution where each grid cell is predictably suitable to the conditions for a species to occur33. To improve model performance and minimize over-fitting, because of the impact of correlation among explanatory variables, a Pearson’s correlation coefficient test was performed for each pair of quantitative variables34–36. MaxEnt25 is a model for predicting the distribution of species based on environmental parameters and species’ presence data37. It uses both continuous and categorical data25. MaxEnt has been reported to be comparable to other highly accurate prediction methods, even when dealing with small sample sizes38. This model was run for the current and future (2050–2070) distributions. MaxEnt model was evaluated through the use of the area under the curve (AUC)39. While generating response curves, the MaxEnt model estimates the relative effect of each predictor40.
Models for the three species performed better than the random, with an AUC values 0.97 (C. procera), 0.98 (P. cinararia), and 0.97 (Ziziphus spina-christi). The jackknife test of variable importance for C. procera concede that out of the 10 selected environmental variables for model execution, precipitation of the coldest quarter (36.3%) temperature annual range (27.2%) and annual precipitation (6.3%) exhibited the highest gain towards the habitat distribution (Figure 1a). Whereas for P. cinararia precipitation of the coldest quarter followed by temperature seasonality and mean diurnal range were the highly influencing variables for its distribution (Figure 1b). At the same time for Ziziphus spina-christi precipitation of wettest month (56.1%) influenced the most followed by precipitation of the coldest quarter and annual precipitation (Figure 1c). Locations for each species are available as Underlying data41.
Results from the current habitat distribution map (Figure 2) clearly indicate that the environmental conditions in the northern part of the UAE (Ras AlKhaimah, Al-Fujayrah, Sharjah, Ajman, Umm Al-Quwain, Dubai and Al-Ain) act as the most suitable habitat for the selected three species. The predicted habitat suitability was reclassified into three classes and this unfolded that only 1.9% of the country showed high habitat suitability (H) for C. procera (Figure 3) followed by 9.4% as medium suitability habitat (M). The remaining 88.6% of the country has low environmental suitability (L) for this particular species. For P. cinararia (Figure 3), 1.5% of the total area was predicted as a highly suitable habitat and 5.7% as medium suitability. Ziziphus spinachristi (Figure 3) revealed high and medium suitability habitat of around 5.7% and 11.5%, respectively.
Current and future habitat distribution (current, 2050s and 2070s) for Calotropis procera (a), Prosopis cineraria (b) and Ziziphus spina-christi (c).
The future (2050s) habitat distribution of C. procera indicated a slight reduction in the high (1.6%) and medium (8.8%) suitability classes. As for 2070s, the model predicted a visible increase in the high suitability class to 7% and medium suitability to 11.9%. The northern part of the country remains the highest environmentally suitable habitat for C. procera. For P. cinararia it is predicted that by 2050s, high suitability will be 8.5% and medium suitability will be 12.3%. In contrast, and by 2070s the high- and medium-suitability classes will see a reduction to 7% and 10.3% of the total area, respectively. For Ziziphus spina-christi distribution, an increase of its high habitat suitability distribution for both 2050s and 2070s, as climate is expected to change.
In this study we predicted the suitable habitat for three significant dry land species with the MaxEnt model. Both current and future habitat distribution models of the selected three species were efficiently projected by MaxEnt. Both C. procera and P. cinararia had less area in the high habitat suitability for the present scenario. Whereas Ziziphus spina-christi had comparatively high present suitable habitat in the country. All three species were projected to have an increased level of high habitat suitability in the future. Calotropis procera can easily adapt with harsh climate conditions especially they rapidly adjust with water availability and loss42. A study by Frosi et al.43 suggested that C. procera has high water use efficiency because of its high photosynthetic rate, despite a reduced stomatal conductance, which acts as its fundamental strategy to survive harsh growth conditions. P. cineraria is the national tree of UAE and most of the artificial forest and road sides in the country are afforested by this plant. It normally grows on soils ranging from sandy to clay loam Kumawat44.
The present study can be useful for selecting species like p. cinararia for forest and roadside plantation in the future floral conservation programs. Predicted areas for P. cineraria consist of a large area of artificial forest27. In UAE, during the past few decades around 300,000 ha has been planted with forests45. The main purpose of these plantation is to combat desertification by afforestation. These artificial forests utilizes numerous resources with no economic benefit.
Other than P. cineraria, these forests consists of many other native and invasive species including P. juliflora, Salvadora persica and Ziziphus spina-christi. P. cineraria can have a positive impact on specie diversity and density27. Prosopis cineraria improves organic matter, soil physical conditions and water-holding capacity. Therefore, it outperforms other species in the same area. Soil nutrient content is higher beneath the P. cineraria trees27. Growth and floral diversity are better under P. cineraria than on adjacent areas46.
Ziziphus spina-christi can resist heat and drought and its deep taproot provide an extraordinary regenerative potentials. This might be the reason for larger area distribution of this species in the UAE. Yet these species’ future habitat suitability areas are important for the country to incorporate in the conservation strategies under climate change. The areas predicted to be suitable can be considered for afforestation programs in the UAE. For all the three species, and with climate change impact, maximum habitat suitability predictions will be in the north-eastern parts of the UAE. It is important to highlight that one of the most vulnerable areas to climate change are dry land ecosystems47.
The MaxEnt model provides useful information about species and their satiable habitats, which can be used as a basis in planning future conservation and afforestation programs. Additionally, the model is also efficient in predicting species distribution under different climate change scenarios. Conservation and afforestation programs considering the species and their suitable habitats, therefore, will not only be easily manageable and sustainable but also economically efficient by using different SDMs. Finer spatial scales of species distribution modeling may be an appropriate next step toward species conservation under the potential changes in climatic conditions.
Figshare: SURE2018.zip. https://doi.org/10.6084/m9.figshare.8233328.v141. This project contains a zipped folder made up of the following files:
Calotropis procera.csv (location of Calotropis procera this study).
Prosopis cineraria.csv (location of Prosopis cineraria in this study).
Ziziphus spinachristi.csv (location of Ziziphus spinachristi in this study).
Data are available under the terms of the Creative Commons Attribution 4.0 International license (CC-BY 4.0).
This project was funded by UAE University (Project: 31S347) as part of the Summer Undergraduate Research Experience (SURE+).
The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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