A
Ecological Drivers of Cogon Grass (Imperata Cylindrica) Invasion: A Global Assessment of Range Expansion under Wildfire and Climate Change
Pradeep Adhikari 1
Yong Ho Lee 1,2
James H. Thorne 3
Prabhat Adhikari 4
Anil Poudel 4
EunSeo-Lee 5
Dean R. Brookes 6
Pun Maya Maharjan 7
Sun Hee Hong 4,8✉ Phone+82-41-950-5532 Email
Changwan Seo 5✉ Email Email
1 Institute of Humanities and Ecology Consensus Resilience Lab Hankyong National University 17579 Anseong Republic of Korea
2 OJEong Resilience Institute Korea University 02841 Seoul Republic of Korea
3 Department of Environmental Science and Policy University of California 95616 Davis United States of America
4 School of Plant Science and Landscape Architecture, College of Agriculture and Life Sciences Hankyong National University 17579 Anseong Republic of Korea
5
A
National Institute of Ecology 33657 Seocheon Republic of Korea
6 United States Department of Agriculture, Agricultural Research Service, Australian Biological Control Laboratory CSIRO Health and Biosecurity GPO Box 2583 4001 Brisbane Queensland Australia
7 G+FLAS Life Sciences 123 Uiryodanji-gil, Osong-eup, Heungdeok-gu 28161 Cheongju-si Korea
8 Hankyong National University 17579 Anseong Republic of Korea
9
A
A
+82-31-670-5087
Pradeep Adhikaria†, Yong Ho Leea,b†, James H. Thornec, Prabhat Adhikarid, Anil Poudeld EunSeo-Leee, Dean R. Brookesf, Pun Maya Maharjang Sun He Hongd,*, Changwan Seoe,*
aInstitute of Humanities and Ecology Consensus Resilience Lab, Hankyong National University, Anseong 17579, Republic of Korea.
bOJEong Resilience Institute, Korea University, Seoul 02841, Republic of Korea.
cDepartment of Environmental Science and Policy, University of California, Davis, 95616, United States of America.
dSchool of Plant Science and Landscape Architecture, College of Agriculture and Life Sciences, Hankyong National University, Anseong 17579, Republic of Korea.
eNational Institute of Ecology, Seocheon 33657, Republic of Korea.
fUnited States Department of Agriculture, Agricultural Research Service, Australian Biological Control Laboratory, c/o CSIRO Health and Biosecurity, GPO Box 2583, Brisbane, Queensland 4001, Australia.
g G + FLAS Life Sciences, 123 Uiryodanji-gil, Osong-eup, Heungdeok-gu, Cheongju-si 28161, Korea
* Corresponding authors
Changwan Seo, E-mail: dharmascw@gmail.com; dharmascw@nie.re.kr
National Institute of Ecology, Seocheon 33657, Republic of Korea
Phone: +82-41-950-5532
Sun Hee Hong, E-mail: shhong@hknu.ac.kr
Hankyong National University, Anseong 17579, Republic of Korea
† These authors contributed equally to this work.
Phone: +82-31-670-5087
Running title
Global risk assessment of cogon grass
Abstract
A
Assessing habitat suitability and potential spread of invasive species is vital for identifying vulnerable regions and guiding management actions to protect biodiversity and ecosystem services. We evaluated the global habitat suitability of cogon grass (Imperata cylindrica), one of the world’s 100 worst invasive species, using species distribution modeling with five machine learning algorithms. Shared Socioeconomic Pathway (SSP) scenarios integrating climate, soil, wildfire regimes, and human population dynamics were used to determine ecological drivers and invasion risks. Wildfire emerged as a key factor promoting cogon grass expansion. Globally, suitable habitat is projected to increase by 20.3%, 40.3%, 42.8%, and 44.4% under SSP1-2.6, SSP2-4.5, SSP3-7.0, and SSP5-8.5, respectively. Currently, 41 tropical and subtropical countries already exhibit high habitat suitability, with cogon grass established across grasslands, savannas, steppes, and alpine meadows. Under SSP5-8.5, 70 countries—particularly in Africa and South America (e.g., Angola, Congo, Haiti, Trinidad and Tobago)—are projected to shift to highly suitable conditions by 2061–2080. The Asian alpine and subtropical regions, along with African and South American savannas, are especially vulnerable. These findings reveal significant global invasion risks under future environmental change and provide essential insights for strengthening biosecurity, early detection, and sustainable management strategies.
Keywords:
cogon grass
climate change
global scale
fire
habitat suitability
MaxEnt
SSP scenarios
Introduction
Cogon grass (Imperata cylindrica) is a rhizomatous perennial grass that grows in a wide range of habitats, including grasslands, degraded forests, and disturbed areas 13. Its taxonomy is complex, with several recognized varieties, including I. cylindrica var. major; Africana; europaea; condensate; and var. koenigii. These varieties exhibit native distribution ranges across South Asia, Southeast Asia, Africa, the Mediterranean region, South America, and Japan (Fig. 1) 1,4. This species thrives at elevations ranging from 0–2,700 m above sea level and in temperate to tropical climates, with mean annual temperature and precipitation levels ranging from 15–30°C and 50–500 cm, respectively 5.
Fig. 1
Global occurrence records of cogon grass (n = 2,113) downloaded from the Global Biodiversity Information Facility (GBIF) database (GBIF, 2024). The green points indicate the native range of cogon grass, and the red points indicate the introduced points of cogon grass. The map was generated using ArcGIS Desktop 10.8 (https://desktop.arcgis.com).
Click here to Correct
Cogon grass is one of the world's 100 worst invasive weeds, and its invasive nature is attributed to traits that enable adaptation in poor soils, tolerance to prolonged heat waves, heavy rainfall, and excessive drought, and genetic plasticity 5,6. Cogon grass exhibits monotypic habitat expansion through an extensive rhizome network and produces prolific seeds that are dispersed by wind over long distances, allowing it to colonize uninfested land 79. Moreover, cogon grass can produce phytotoxic compounds and allelopathic chemicals in soil that interfere with the physiological processes of native plants and disrupt the soil microbial community 2.
Global climate change, including increasing temperatures and shifts in precipitation patterns, is projected to increase the invasion success of cogon grass 10. As a C4 plant with high resilience to various climatic conditions, cogon grass possesses competitive advantages over C3 plants in terms of absorbing nutrients and water, which may amplify its expansion into newly suitable habitats5,9. One key factor influencing its spread is fire occurrence, a natural disturbance that plays a significant role in shaping the invasion success of several alien and invasive plant species. Climate change further exacerbates this issue by contributing to the occurrence of more frequent and intense wildfires through altered weather patterns, increased temperatures, and prolonged droughts 11; conditions that increase the invasion success of many alien species, including cogon grass8. Cogon grass’s underground rhizomes constitute 60% of its biomass and are well protected from heat, enabling rapid regeneration after fires1. Additionally, its dry, flammable leaves and stems promote recurring intense fires, for which this species exhibits tolerance, allowing it to quickly colonize disturbed areas7,12.
The intentional introduction of cogon grass into nonnative regions, including the United States, began in the 19th century for purposes such as forage crop cultivation, soil erosion control, ornamental purposes (e.g., Japanese blood grass), pulp production, and the extraction of phytochemicals for pharmacological and industrial uses4,10,13. Currently, this invasive species has been deliberately introduced into more than 73 countries and now invades approximately 500 million hectares globally, including over 100,000 hectares in the United States, including Florida, Alabama, and Mississippi states 9,14. Its spread has significant economic, environmental, and social consequences worldwide. In Asia and Africa, the invasion of cogon grass affects more than 35 crops, with estimated decreases in production by 40% in Southeast Asia and 62–80% in West Africa for key crops such as maize, cassava, rubber, and tea1,10. Similarly, in the United States, it imposes substantial economic losses, costing Florida alone an estimated $30.2 million annually 15.
Understanding the habitat suitability of cogon grass and its ecological drivers—such as global climate and land cover changes, along with the exacerbation of invasion by wildfires—is crucial for predicting current and future distributions, identifying vulnerable regions, and evaluating its impacts on natural and agricultural biodiversity16. Additionally, such studies enable the identification of proactive management and mitigation strategies, including robust quarantine protocols, early detection, and cogon grass eradication measures6.
Within this context, species distribution models (SDMs) are useful tools for predicting the spread of invasive weeds via spatial data on climate and human disturbances17. SDMs allow cost-effective and time-efficient evaluations of potential invasion risks in combination with invasion history and environmental suitability18,19. Globally, SDMs are increasingly employed to predict the habitat suitability of alien and invasive species in both aquatic and terrestrial ecosystems at various levels from local to global levels19. In our earlier studies, we performed SDM-based global risk assessments of many world’s worst invasive species, such as Ardisia elliptica, Lantana camara, Myocastor coypus, and Oxalis latifolia 18,2022. These assessments identified high-risk countries under current climatic conditions, and new introductions and substantial increases in invasion risks in several countries worldwide in the future were predicted16.
The incorporation of bioclimatic variables along with other environmental variables, including land use and land cover changes, biome factors, soil conditions, human footprints, and wildfires, can significantly increase the prediction accuracy of SDMs16,23,24. Therefore, in this study, we integrated climatic variables with wildfire occurrence and land cover change in the modeling of cogon grass range dynamics with the following objectives: (1) to assess model performance with and without the inclusion of wildfire and land cover change variables; (2) to project current and future spatial distributions of cogon grass worldwide; (3) to explore the relationship between changes in climatic variables and habitat suitability worldwide; and (4) to evaluate habitat suitability and categorize countries into distinct risk levels. This approach offers a comprehensive understanding of the ecological interactions and spatial variations influencing species distributions, thereby contributing to the formulation of effective management strategies for invasive species.
Results
Model evaluation and selection of the optimal model
Two sets of input variables, including environmental variables with and without wildfire and environmental variables, were employed with five machine learning algorithms to predict the spatial distribution of cogon grass. The MaxEnt model with environmental variables and wildfires as input variables demonstrated the best prediction performance among the five algorithms, with an AUC value of 0.92, a TSS value of 0.73, a kappa value of 0.68, a sensitivity value of 0.79, and a specificity value of 0.81 (Table 2). Consequently, the outputs of the MaxEnt model were used to predict the spatial distribution of cogon grass and the habitat suitability of this species worldwide. Considering the values of the chosen evaluation metrics, the results indicated excellent prediction performance and strong agreement between the observations and predictions. The results also suggested that wildfire occurrence is the most significant ecological factor influencing the expansion of cogon grass habitats.
Table 2
Comparison of predictive performances of five SDMs based on the model evaluation parameters
Model+
With wild fire ++
Without wild fire +++
name
AUC
TSS
Kapp
Sens
Spec
AUC
TSS
Kapp
Sen
Spe
BRT
0.76
0.54
0.44
0.69
0.68
0.75
0.53
0.42
0.61
0.53
GAM
0.72
0.69
0.55
0.71
0.72
0.71
0.61
0.4
0.65
0.62
RF
0.85
0.72
0.55
0.72
0.74
0.8
0.65
0.55
0.70
0.72
MaxEnt
0.92
0.73
0.68
0.79
0.81
0.89
0.71
0.67
0.71
0.77
XGB
0.72
0.72
0.63
0.67
0.68
0.72
0.68
0.64
0.64
0.56
+, The column shows five machine learning models: BRT (boosted regression trees), GAM (generalized additive model), RF (random forest), MaxEnt (maximum entropy), and XGB (extreme gradient boosting). ++, Modeling variables included six bio climatic variables e.g., annual mean temperature, mean diurnal range, isothermality, annual precipitation, precipitation in the wettest month, and precipitation in the driest month, three soil variables e.g., soil carbon, soil moisture, and soil pH, human influence index, and wildfire. +++, Modeling variables included bioclimatic variables, soil variables, and human influence index. The abbreviations AUC, TSS, Sens, and Spe represent the area under the receiver operating characteristic curve, true skill statistic, sensitivity, and specificity, respectively.
Evaluation of the environmental variables
A
Pearson's correlation analysis was conducted among the 19 bioclimatic variables and five environmental variables. On the basis of the lower Pearson correlation coefficient values (r ≤ 0.75), three temperature-related variables (Bio01, Bio02, and Bio03), three precipitation-related variables (Bio12, Bio13, and Bio14), and six additional environmental variables (wildfires, HII, land use and land cover change, soil carbon, soil moisture, and soil pH) were selected for cogon grass modeling (Table S2). Among these 11 modeling variables, wildfires contributed the most to the model outcomes, at 27.14%, under the current and future climate change scenarios (SSP1-2.6, SSP2-4.5, SSP3-6.0, and SSP5-8.5) from 2061–2080. Similarly, the temperature-related variables Bio1 and Bio3 and the precipitation-related variable Bio12 yielded nearly equal contributions to the model outcomes, with average relative contributions of 20.76%, 20.23%, and 19.51%, respectively (Table 1). These findings suggest that wildfire occurrence, annual mean temperature, isothermality, and annual precipitation are four critical environmental determinants influencing cogon grass habitat expansion and invasion of new ecosystems. The other variables play a minor role in the model.
Table 1
Contribution of environmental variables in MaxEnt modeling of I. cylindrica
Code
Description
Unit
Model contribution (%)+
Bio1
Annual mean temperature
°C
19.76
Bio2
Mean diurnal temperature range
°C
1.29
Bio3
Isothermality (BIO2/BIO7) (×100)
%
17.23
Bio12
Annual precipitation
mm
19.51
Bio13
Precipitation in the wettest month
mm
0.04
Bio14
Precipitation in the driest month
mm
6.73
HII
Human influence index
-
0.53
Land
Land cover change
-
5.25
Fire
Wild fire
-
27.14
Soil C
Soil carbon
-
0.05
Soil M
Soil moisture
-
2.08
Soil pH
Soil pH
-
0.31
+, Average contribution in MaxEnt model estimated under the current and future climate change scenarios SSP1-2.6, SSP2-4.5, SSP3-6.0, and SSP5-8.5 by 2060 − 2080.
Influence of wildfire on the expansion of cogon grass habitats
A
A
The global habitat suitability of cogon grass was predicted on the basis of environmental variables, with and without wildfires, and current suitable areas worldwide were identified (Fig. 2A). Suitable areas were predicted on all continental regions regardless of the inclusion of wildfires in the model. However, the proportion of the suitable area was greater on each continent when wildfires were included as an environmental variable (Fig. 2A). The expansion of suitable area from current extents increased by up to 23.1% globally, with the highest increase observed in Africa (65.5%) compared with the model without wildfire (Table S3). Several countries in Asia (e.g., Nepal, India, and Indonesia), Oceania, Africa (e.g., Ghana, Ivory Coast, the Central African Republic, and Nigeria), South America (e.g., Brazil, Bolivia, Paraguay, and Venezuela), Europe (e.g., Slovenia, Croatia, and Greece), North America (e.g., the United States) and Australia exhibited habitat expansion when wildfire was considered (Table S4 and Fig. S2). Expanded range was located primarily in tropical and temperate grassland biomes, including the African savanna, Australian savanna, and Campos. These results suggest that wildfire occurrence is a significant factor contributing to the expansion of cogon grass habitats.
Fig. 2
Potential global distribution of cogon grass under the current environmental conditions. Fig. A shows the modeling results when global species occurrence points and environmental variables are used, with and without wildfires. The red, blue, and yellow colors denote the predicted areas based on the environmental variables with wildfires, without wildfires, and the overlap of both, respectively. Similarly, the green, and red, colors in Fig. B denote the predicted distributions of cogon grass based on only existing occurrence points in native regions and invading regions (North and South America), respectively. Similarly, the yellow color indicates predicted areas overlap estimated between both. The map was generated using ArcGIS Desktop 10.8 (https://desktop.arcgis.com).
Click here to Correct
Habitat suitability of cogon grass under current environmental conditions
The global spatial distribution of cogon grass under the current environmental conditions was predicted via species distribution modeling with the MaxEnt algorithm (Fig. 2A). Under the current conditions, suitable areas for cogon grass in nonnative regions are primarily concentrated along the western coast of Central Africa, southeastern region of Africa, southeastern United States and the Caribbean region, southern Europe, and central part of South America, approximately between 40° north and 35° south latitudes. The current global suitable area is estimated to 27,008,598 km2 representing approximately 14.2% of the Earth's total land surface area (Table 3). Among the global continental regions, Oceania, South America, and Africa exhibited relatively high proportions of suitable areas, covering 30%, 27%, and 16.6% of their respective land areas. In contrast, other continents, including Asia, Europe, North America, and Antarctica, exhibited lower proportions of suitable areas, covering less than 13.5% of the total land area (Table 3).
Table 3
Proportion of change in suitable habitats of I. cylindrica in different continents under the global climate change
Continents
Total
Area (Km
) +
Current
Future climate change scenarios ++
SSP1-2.6
SSP2-4.5
SSP3-7.0
SSP5-8.5
North America
45,923,957
3,455,824
9.12
14.74
14.29
10.54
Asia
65,097,657
8,769,872
11.89
28.05
27.83
27.67
Europe
18,937,544
1,680,290
12.36
14.08
17.92
17.60
Africa
31,675,434
5,245,613
44.84
80.26
104.77
124.77
South America
18,992,283
5,135,168
24.45
57.08
44.12
44.34
Oceania
9,017,896
2,721,884
12.25
19.55
20.34
3.06
+ Total area of each continent is estimated based on the number of cells count in each continent and area of each cell at 2.5 min resolution is approximately 21.5 km2 at equator. ++ Change in the proportion of suitable habitats of cogon grass under different climate change scenarios—Shared Socioeconomic Pathways (SSP1-2.6, SSP2-4.5, SSP3-7.0, and SSP5-8.5)—by 2061–2080 compared to the current suitable habitat. The Oceania region includes continental mass of Australia, New Zealand and other Oceanic Islands.
A
Similarly, two sets of species occurrence points, representing only native range regions and introduced regions (North and South America), along with the same set of environmental variables, including wildfires, were selected and modeled separately. The results revealed that potentially suitable areas overlapped across all continents (Fig. 2B), with the highest proportion in South America (18%), followed by Oceania (14.4%), North America (6.6%), Asia (4.6%), Europe (1.7%), and Africa (0.9%) (Table S5). North America, particularly the United States, exhibited high invasion by cogon grass on the basis of the prediction using only introduced points.
The extent of suitable range obtained from introduced points was greater than that obtained from native points in certain countries within the introduced regions, e.g., the United States and Mexico (Fig. 2B), suggesting that invasive species conserve much of their original niche in the new environment and can thrive wherever suitable conditions exist, regardless of geographic origin.
Suitable range of cogon grass under future environmental conditions
The global distribution of cogon grass was estimated under the SSP1-2.6, SSP2-4.5, SSP3-6.0, and SSP5-8.5 climate change scenarios. We measured the expansion of suitable areas between 2061 and 2080 (Fig. 3) and found an increase in habitat suitability for cogon grass on all continents. The total area of suitable habitats was projected to increase, with estimated increases of approximately 20.4%, 40.3%, 42.8%, and 44.4% under the SSP1-2.6, SSP2-4.5, SSP3-7.0, and SSP5-8.5 scenarios, respectively, relative to the current habitat suitability of cogon grass.
Fig. 3
Global spatial distribution of cogon grass under the current and future environmental conditions. The maps show the global spatial distributions of cogon grass under the current conditions and future environmental scenarios, including climate change SSP1-2.6 (A), SSP2-4.5 (B), SSP3-7.0 (C), and SSP5-8.5 (D) scenarios, as well as the influences of wildfires, soil properties, and the human influence index for the 2061–2080 period. The green areas denote the current potential habitats, and the red areas denote the projected habitat expansions from 2061–2080.
Click here to Correct
Asia, Africa, and South America were predicted to exhibit relatively high rates of habitat expansion, with estimated increases ranging from 11.9–44.8% (SSP1-2.6), 28.1–80.3% (SSP2-4.5), 27.8–104.8% (SSP3-6.0), and 27.7–124.8% (SSP5-8.5) compared with the current extents of suitable areas (Table 3). North America and Europe, both introduced regions of cogon grass, are likely to face significant impacts in their southeastern and southern areas, respectively, with an estimated 14.3% and 17.9% of their respective land surfaces affected under the SSP3-7.0 (Table 3). These findings suggest that the temperature and precipitation patterns under the SSP3-7.0 scenario could occur within a favorable range for the distribution of cogon grass. Overall, the results indicated that global warming is likely to increase suitable environments worldwide for expansion of cogon grass.
Changes in habitat suitability extent by country under future environmental changes
Cogon grass currently occurs in 73 countries worldwide, of which > 50 countries in Asia, Africa, Europe, and Oceania are part of its native range, whereas the remaining countries fall within its introduced range. The mean area of suitable habitat of cogon grass was estimated across 195 countries and territories globally and classified into five categories under current and future climate change scenarios (SSP1-2.6, SSP2-4.5, SSP3-7.0, and SSP5-8.5) (Fig. S2). Currently, 36 countries, such as Belgium, Mongolia, Rwanda, and Iceland, encompass climatically unsuitable habitats for cogon grass (Fig. 4). Similarly, 70 countries, particularly in the northern parts of Asia, Africa, and Europe, were estimated to exhibit poor habitat suitability, whereas 41 countries located in tropical and subtropical regions were found to exhibit extreme habitat suitability (Table S4).
Fig. 4
Estimated number of countries classified into different categories of habitat suitability for cogon grass in different countries and territories under the current climate and future climate change scenarios, namely, SSP1-2.6, SSP2-4.5, SP3-7.0, and SSP5-8.5, from 2061–2080. Habitat suitability was classified into five categories, i.e., unsuitable (0), poorly suitable (0.01–0.25 area of the country), moderately suitable (0.25–0. 5), highly suitable (0.5–0.75), and extremely suitable (0.75–1.0), across 195 countries worldwide. The different colors in the pie chart indicate the number of countries within each habitat suitability category. The map was generated using ArcGIS Desktop 10.8 (https://desktop.arcgis.com).
Click here to Correct
Global climate change, fires, and human disturbances are major factors contributing to the expansion of the habitat of cogon grass. Under two high-emission scenarios, namely, SSP3-7.0 and SSP5-8.5, this species is projected to encounter extremely suitable areas in 68 and 70 countries, respectively, from 2061–2080 (Fig. 4). Furthermore, several countries currently classified as unsuitable or poorly suitable are predicted to shift into the highly suitable or extremely suitable category in the future, with these areas estimated to cover more than 50% of the land surface of each country. Under the SSP1-2.6, SSP2-4.5, SSP3-7.0, and SSP5-8.5 scenarios, four, 14, 16, and 17 countries are respectively projected to become highly invasable (Table 4). We found that the rate of range expansion is greater in African and Caribbean regions than in the currently suitable areas. These modeling outputs and areas of suitable habitats underscore the highly invasive nature of cogon grass and suggest that future environmental changes will increase its invasion area up to 44.4% (SSP5-8.5). If efforts to control greenhouse gas emissions are not effective, at least 14 countries could face serious impacts on their natural and agricultural ecosystems.
Table 4
Change in habitat suitability of cogon grass in different countries of the world under the future climate change scenarios
Transition of habitat suitability of cogon grass under the global climate change scenarios +
SSP1-2.6
SSP2-4.5
SSP3-7.0
SSP5-8.5
Burundi
Dominica
Kiribati
Marshall Islands
Angola
Burundi
Guyana
Haiti
Central African Republic
Equatorial
Guinea
Indonesia
Kiribati
Marshall Islands
Malaysia
Suriname
Trinidad and Tobago
Zaire
Angola
Burundi
Cape Verde
Central African Republic
Equatorial Guinea
Guyana
Haiti
Indonesia
Kiribati
Malaysia
Marshall Islands
Rwanda
Sierra Leone
Suriname
Trinidad and Tobago
Zaire
Angola
Burundi
Cape Verde
Central African Republic
Congo
Equatorial Guinea
Guyana
Haiti
Indonesia
Kiribati
Malaysia
Marshall Islands
Rwanda
Sierra Leone
Suriname
Trinidad and Tobago
Zaire
+, Countries that currently have unsuitable (0) or poorly suitable (0.01–0.25) habitats are projected to change into highly suitable (0.5–0.75) or extremely suitable (0.75–1.0) habitats by 2061–2080 under the climate change scenarios SSP1-2.6, SSP2-4.5, SSP3-7.0 and SSP5-8.5.
Habitat coverage of cogon grass in different land cover types
Suitable conditions for cogon grass were analyzed across various global land cover types, revealing a projected significant expansion under future environmental change. Currently, the highest proportion of cogon grass coverage is observed in artificial lands including human settlements, fragmented landscapes, and industrial areas (42.5% of these lands globally), followed by agricultural land (35.3%) and mixed land cover types (24.7%) (Table 6). Under the SSP5-8.5 scenario, coverage is expected to increase to 51.3% in artificial areas, 44.7% in agricultural land, 28.1% in forests, and 19.4% in grasslands, indicating growing ecological threats to native biodiversity and ecosystem stability. Wetlands and water bodies, including marshy and poorly drained soils, also show a steady increase from 12.4% to 17.7%, suggesting cogon grass’s strong adaptability to moist environments. Although non-vegetated areas maintain very low coverage, a slight increase from 0.1% to 0.3% reflects its minor expansion potential in harsh conditions. Overall, these trends highlight the ecological plasticity of cogon grass and its increasing potential to invade diverse landscapes under climate change.
Table 6
Global proportion of area covered by cogon grass in each category of land cover under the current and future climate change scenarios
Land cover
Total area (Km
)+
Current
SSP1-2.6
SSP2-4.5
SSP3-7.0
SSP5-8.5
Artificial
5,946,793
42.49
46.82
49.99
50.68
51.32
Mixed
21,478,238.4
24.70
28.08
31.03
31.12
31.62
Wetlands
2,714,889.6
12.38
14.35
17.56
17.59
17.70
Agriculture
14,515,598.6
35.29
40.58
43.90
44.23
44.73
Forest
50,463,917.8
16.33
20.20
25.71
26.18
28.06
Grassland
57,582,007.2
13.27
16.44
18.74
19.35
19.38
Non-vegetated
40,175,632.4
0.11
0.20
0.25
0.26
0.29
+, The area of each category of land use and land cover change is estimated based on the number of cells count in each category of land and area of each cell at 2.5 min resolution is approximately 21.5 km2 at equator. The artificial category includes disturbed areas such as human settlements, land fragmentation, and infrastructure development.
Habitat coverage of cogon grass in grasslands of different continents and Savanna Regions
A
Cogon grass currently overlaps with major grassland regions on all six global regions, and this overlap it is projected to expand on grasslands under all future climate change scenarios (Fig. 5). Presently, cogon grass is found in five types of grassland in Asia, 17 in Africa, 18 in Australia, 10 in North America, 11 in South America, one in Europe, and two in Oceania (Table S6). These ecosystems represent a wide variety of grassland formation types, including alpine meadows, temperate steppes, tropical savannas, and semi-desert scrublands—each shaped by distinct climatic and topographic conditions.
Fig. 5
Spatial distribution of cogon grass in global grasslands under the current (A) and future climate change scenarios SSP1-2.6 (A), SSP2-4.5 (B), SSP3-7.0 (C), and SSP5-8.5 (D) for the period 2061–2080. The green color in maps shows area of global grasslands, blue color indicates habitat distribution of cogon grass, and purple color indicates habitat overlap in global grasslands. The map was generated using ArcGIS Desktop 10.8 (https://desktop.arcgis.com).
Click here to Correct
Model projections under future climate scenarios indicate a notable expansion of cogon grass habitat, both within grassland types it currently occupies and into neighboring ones. This trend is particularly evident under high-emission scenarios SSP3-7.0 and SSP5-8.5. For instance, in the “Eastern Himalayan alpine shrub and meadows” ecoregion—classified within the Central Asian Alpine Scrub, Forb Meadow & Grassland formation—and the “Campos Rupestres montane savanna” ecoregion—part of the Brazilian-Parana Montane Shrubland and Grassland formation—cogon grass is projected not only to persist but also to expand substantially under all scenarios from SSP1-2.6 through SSP5-8.5 (Table S6).
A
Range area of cogon grass was further estimated in savanna regions across three continents—Africa, Australia, and South America—under both current and future climate scenarios. At present, cogon grass occupies approximately 1,954,178 km2 in the African Savanna, 1,293,612 km² in the Australian Savanna, and 1,251,408 km2 in the South American Savanna (Table S7). Under the SSP5-8.5 scenario, range area is projected to increase by 58.3% in the African Savanna and 36.5% in the South American Savanna. In contrast, the Australian Savanna is expected to experience only a modest increase of 1.7%. These results suggest that global climate and environmental changes are likely to facilitate the expansion of cogon grass, particularly within major grassland ecosystems, with the most pronounced effects observed in tropical savanna regions.
Discussion
Modeling habitat suitability of cogon grass
Our analysis of the distribution models and environmental variables, revealed that the MaxEnt model with environmental variables and wildfire provided the most accurate projection of the current and potential future ranges of cogon grass (Table 2), and that wildfire is the greatest contributing ecological driver for determining the habitat suitability of cogon grass (Table 1). We found that 41 countries and 70 countries currently contain extremely suitable for cogon grass (Fig. S2). At least 17 countries, including Angola, Congo, and Suriname, which currently present unsuitable or poorly suitable, are projected to shift to extremely suitable conditions in the future under the SSP5-8.5 scenario (Table 5). Analysis of cogon grass distribution across different land cover categories shows that the species currently is most prevalent in disturbed areas such as urban, semi urban, and agricultural areas (Table 6). The high habitat suitability of cogon grass in artificial and agricultural lands arises because these environments reduce competition and provide open niches. The grass’s fire resilience, extensive rhizome network, efficient resource use, allelopathy, and rapid colonization enable it to dominate and persist. Under the SSP5-8.5 scenario, suitable conditions for cogon grass are projected to expand further within these land cover classes and also into forests and grasslands, underscoring the growing threat it poses to native biodiversity and ecosystem stability. Furthermore, cogon grass is found across a wide range of grassland ecosystems, including alpine meadows, temperate steppes, tropical savannas, and semi-desert scrublands globally. This wide ecological range increases the risk to various grassland types. In particular, regions such as the Asian alpine and subtropical zones, as well as the savannas of Africa and South America, are especially vulnerable to future invasions.
Table 5
List of countries having high rate of wild fire estimated between 2012 and 2024.
S. No.
Average annual wildfire +
Average burn area ++
1
Angola
Madagascar
Angola
Myanmar
2
Argentina
Mali
Belarus
Nigeria
3
Belarus
Mexico
Belize
Paraguay
4
Bolivia
Mozambique
Benin
Senegal
5
Brazil
Myanmar
Bolivia
Sierra Leone
6
Cameroon
Nigeria
Cameroon
South Sudan
7
Central African Republic
Paraguay
Central African Republic
Tanzania
8
Chad
Russia
Cote d'Ivoire
Togo
9
China
Senegal
Democratic Republic of Congo
Uganda
10
Colombia
Sierra Leone
Gambia
Venezuela
11
Congo
South Africa
Ghana
Zambia
12
Cote d'Ivoire
South Sudan
Guatemala
 
13
Democratic Republic of Congo
Sudan
Guinea
 
14
Ethiopia
Thailand
Guinea-Bissau
 
15
Ghana
Uganda
Honduras
 
16
Guinea
United States
Laos
 
17
India
Venezuela
Liberia
 
18
Indonesia
Zambia
Madagascar
 
19
Kazakhstan
Thailand
Malawi
 
20
Laos
Uganda
Mozambique
 
21
Tanzania
     
+ and ++ indicate the list of countries having average annual wild fire cases above 5,000 and average annual burnt area above 5% of the total land surface of the country estimated between 2012 and 2024 (Global Wildfire Information System, GWIS, 2024).
Weed species like cogon grass are rarely at equilibrium, exhibiting rapid expansion across new environments25. This dynamic behavior makes it critical to use predictive models that capture nonequilibrium patterns. Our MaxEnt model identified key climatic and environmental drivers, including temperature, precipitation, wildfires, and soil characteristics, that determine habitat suitability. Results indicate that cogon grass thrives under warm climates with variable rainfall and frequent disturbances, such as wildfires. These conditions create opportunities for rapid colonization and expansion into previously unsuitable regions, highlighting the species’ potential to alter ecosystem composition and challenge management efforts across diverse habitats.
Wildfire as a driver of cogon grass invasion
Global climate change is increasing the frequency, intensity and duration of wildfires by altering weather patterns, with increasing temperatures and prolonged droughts impacting native vegetation, ecosystem function, and forest resilience11,26. Wildfires are major ecological disturbances that shape the structure and composition of many ecosystems, and play a crucial role in shaping plant invasions by serving both as a regulator of natural ecosystems and a driver of pyrogenic succession for alien plant species 27.
Alien and invasive species are particularly opportunistic in utilizing postfire habitats because of the presence of open canopies, a reduced number of competitive native species, and improved soil conditions 8. Numerous studies have revealed the rapid invasion of alien plant species following wildfires in both grassland and forest ecosystems 11,28. This phenomenon can lead to landscape transformation and a reduction in the abundance and diversity of native species.
Cogon grass exhibits both fire resistance and fire-altering traits, allowing quick regeneration in open spaces of postfire landscapes 7. The species exhibits high flammability, containing three times the fine-fuel load of many native grasses and creating continuous fuel beds, which lead to the occurrence of frequent fires with greater vertical flame heights1,7. This species provides a self-reinforcing cycle of fires that can destroy native seed banks, increase the mortality of native plants, disrupt ecosystem functions and allow it to grow and expand 5.
Previous studies have indicated that wildfires increase the abundance of cogon grass in burned areas across the United States and other parts of the world through various mechanisms. These mechanisms include increased disturbance, changes in soil chemistry and resource availability, and altered competitive dynamics between invasive and native plant species9,29. Cogon grass thrives in disturbed environments, particularly after fire events, which creates conditions favorable for its spread.
The United States exhibits a high risk of wildfires12, which is amplified by the presence of cogon grass. Increasing temperatures and severe drought lead to the presence of dry vegetation and soil, creating highly flammable fuel that, when combined with high winds, exacerbates the potential for wildfires—particularly in the southeastern and western regions of the United States12,30. In early 2025, southern California experienced a devasting wildfire, followed by outbreaks in Texas and many parts of the southeastern United States, which exhibited abnormal fire activity. These wildfires not only cause damage to native biodiversity but also create conditions for long-term ecological shifts driven by increased plant invasions 31.
Cogon grass also contributes to the mortality of juvenile native plants, which has led to a 40–60% decrease in plant species richness and endemic plant populations in the southeastern USA, particularly in longleaf pine woodlands in Mississippi and the Blackwater River State Forest in Florida 7,8. The rate of fuel accumulation following fires has been measured on a broad scale, and over time, fires in sandhill habitats have caused a shift in the composition of species-rich pine savannas to grasslands dominated by the nonnative I. cylindrica 7. Similarly, in Blackwater River State Forest, Florida, USA, areas that were burned or experienced significant biomass removal following a hurricane supported a greater number of cogon grass patches and larger patch sizes 8. Similarly, in northern California, repeated high-severity wildfires have increased the invasion risk by several nonnative species, including cogon grass, which competes for resources and alters ecosystem dynamics 32. Our study revealed that wildfire occurrence could increase the area of suitable habitats for cogon grass by 11% in North America, particularly in the southeastern United States, spanning from the coastal regions of Florida to Mississippi, Kentucky, and North Carolina (Fig. 2A and Table S3).
Projected increase in wildfire activity across various regions of Africa significantly elevates the invasion risk of cogon grass. This phenomenon is particularly evident in areas exhibiting climate change, land cover change, and the occurrence of nonnative vegetation. Recent studies have shown that tropical forests in Central Africa and western Africa, which are traditionally resistant to fires, now demonstrate frequent wildfire activities due to climate change and deforestation 33. Several countries in Africa, e.g., Angola, Mozambique, the Central African Republic, Ghana, Nigeria, and South Sudan, exhibit over 5000 wildfire ignitions annually, and average annual burned areas above 5% of the national land surface area (from 2012 to 2024; Table 5). Global climate change and increased wildfire create more favorable conditions for cogon grass in such countries. Our study revealed that the greatest proportion of change in the habitat suitability of cogon grass occurred in Central and South Africa, and that several countries in at high risk of invasion.
The Congo River Basin in Africa is one of the largest biodiversity hotspots. The riverine forest of the basin holds nearly 10% of the world's forest-based carbon and supports the livelihoods of more than 50 million people 33. Cogon grass can dominate landscapes in the Congo Basin, outcompeting native flora and altering ecosystem dynamics. In South Africa, the invasion of alien plants has been linked to increased fire activity. For example, the Knysna wildfire of 2017 allowed invasion of the Fynbos ecosystem by several invasive species, including cogon grass, in the southwestern and southern parts of South Africa 27,34.
South America is one of the continents with the most frequent wildfires, and most fires are anthropogenic in origin, occurring mostly in tropical regions during the dry season 35. The increase in wildfire frequency across South America can increase the invasion risk of cogon grass. Our model revealed the wildfire-induced invasion risk of cogon grass in many countries in the northern, central and southern parts of South America, e.g., Brazil, Venezuela, Colombia, Bolivia and Argentina. Our wildfire model results indicated an increase in the area of suitable habitats in South America of 28% compared with the estimate without wildfires. Studies on fire ecology and its role in determining the invasion success of alien plants in South America have not yet been published 36. Therefore, these results may be useful for estimating the impacts of wildfires on native biodiversity and invasive species on a broad scale in South America.
Direct climate effects on cogon grass distribution
Global climate change, through fluctuations in the annual temperature and altered precipitation patterns, can weaken metabolic processes, reduce flowering and seed production, and decrease the growth and productivity of native plants, thereby reducing their competitive ability against invasive species 37,38. These climatic changes tend to favor invasive species, as they generally possess wider physiological tolerance ranges, extended phenologies, and greater competitive abilities, allowing them to establish and spread in new ecosystems 39. Similarly, isothermality influences plant invasion dynamics by regulating temperature stability and the overlap of climatic niches between invasive and native species. Native species typically prefer stable thermal environments, whereas invasive species are often better suited to cope with daily temperature fluctuations 40. As a result, lower isothermality creates conditions that align more closely with the climatic requirements of invasive plants, thus promoting their successful establishment and competitive advantage 25,41.
Our study indicated that the habitat suitability of cogon grass spans a wide range of climatic variables, with mean annual temperatures ranging from 2.6–30.4°C, annual precipitation amounts ranging from 31 to 4,784 mm, and isothermality values between 20.12 and 90.05 under the current climatic conditions. These findings suggest that cogon grass can persist across a broad spectrum of future climatic conditions 9. The species exhibits variable cold tolerance depending on its growth stage and environmental conditions. Mature, dormant plants can survive at temperatures as low as -14°C and can tolerate 1–5 days of freezing at -5°C to -14°C. However, prolonged soil freezing exceeding 10–14 days may be lethal, particularly for young or nondormant plants 5. This is why large areas in the northern parts of Asia, Europe, and North America (e.g., Russia, Mongolia, Iceland, Sweden, Norway, and the United Kingdom) currently exhibit unsuitabile or very poor habitat suitability.
Under global climate change scenarios based on SSPs, the Earth's annual mean temperature is projected to increase by 1.8–4.4°C relative to 1850–1900 levels 42. Future precipitation patterns are predicted to change, with increases in some regions that currently exhibit arid climatic conditions and decreases in regions with high precipitation 43. The rate of precipitation is expected to change by -0.2% to + 4.7% under SSP1-2.6 and by 0.9% to 12.9% under SSP5-8.542. With a warming climate, precipitation is projected to increase at high latitudes (e.g., the average precipitation on the Pothwar Plateau in Pakistan is projected to increase by up to 1,528 mm/year) and in tropical oceans but may decrease in subtropical regions 42,43.
Global climate change is projected to expand suitable areas for cogon grass across both native and introduced regions. Many areas in Africa (e.g., Cameroon, the Democratic Republic of the Congo), South America (e.g., Brazil and Peru), Europe (e.g., Kosovo and Greece), and the southeastern United States, which were previously unsuitable, may become highly favorable. Moreover, parts of its native range, including India, Nepal, Indonesia, and Australia, are also projected to experience increased suitability, consistent with climax-based modeling of cogon grass in a recent study44. These shifts highlight the species’ broad climatic tolerance and signal a growing risk to biodiversity and ecosystem stability worldwide, emphasizing the need for proactive management in both native and invaded landscapes.
Land cover and ecoregional vulnerability
Cogon grass has remarkable adaptability allowing it to colonize a broad spectrum of land cover types, including grasslands, open forests, pastures, agricultural fields, wetlands, riparian zones, scrubland, roadsides, and urban areas 10. Our study on land cover-specific projections shows how cogon grass's ecological adaptability and invasion potential is amplified by climate change. The greatest future habitat area increases are in artificial lands and agricultural lands, showing the species’ strong association with human dominated landscapes 10. These increases indicate a growing risk to urban infrastructure and agricultural productivity, particularly in regions already vulnerable to invasive species. Also concerning is the projected expansion into forested and grassland ecosystems, which are critical reservoirs of native biodiversity 8. These changes imply that cogon grass could alter ecological processes and replace native vegetation, thereby destabilizing ecosystem functions.
We found that cogon grass poses significant ecological threats across a wide range of grassland ecosystems, including alpine meadows, temperate steppes, tropical savannas, and semi-desert scrublands. Globally, the species currently occurs in 64 ecoregions, spanning native ranges in Asia, Africa, Australia, and Southern Europe, and non-native regions in the Americas (Table S6). In its native regions, cogon grass is generally well-integrated, while in non-native regions, particularly in North and South America, it exhibits invasive behavior with substantial ecological impacts. In North America, it has invaded at least 10 ecoregions, such as the California Central Valley grasslands, Nebraska Sand Hills mixed grasslands, and Western Gulf coastal grasslands. In South America, invasions are recorded in at least 11 ecoregions, including the Humid Pampas, Guianan savanna, Uruguayan savanna, and Patagonian steppe. Future projections indicate that suitable conditions may expand to as many as 73 ecoregions globally, highlighting the growing risk of range expansion in non-native regions and the potential threat to biodiversity and ecosystem function.
This expansion is particularly concerning in regions such as the Asian alpine and subtropical grasslands, and the savannas of Africa and South America, where cogon grass is likely to alter fire regimes, outcompete native plant species, and degrade habitat quality for native fauna 9. These ecological impacts may trigger shifts in species composition, reduce biodiversity, and disrupt ecosystem processes 7,31.
Anthropogenic drivers of invasion
The HII variable incorporates a range of anthropogenic activities—such as infrastructure development, land use change, and population pressure—that can disrupt natural biogeographic barriers and facilitate the spread of invasive species (Gallardo et al., 2015). Accurately predicting invasive species distributions without incorporating socioeconomic factors is challenging, since these human-driven processes play critical roles in shaping invasion dynamics 45. We incorporated the HII to improve predictions of the global distribution of cogon grass. Notably, under the highest-consumption scenario, i.e., SSP5-8.5, we observed the greatest expansion of highly suitable areas, indicating a strong link between high-intensity human activity and increased invasion risk. However, this risk is closely tied to associated shifts in temperature and precipitation patterns under each SSP scenario. Low-emission scenarios, such as SSP1-2.6, exhibit a reduced overall risk, emphasizing the role of sustainable development pathways in mitigating future invasions.
Management implications and control strategies
The global management of cogon grass presents significant challenges due to its aggressive growth, adaptability, and regenerative capacity 9; and continuous monitoring, stakeholder involvement, and sustained funding are critical for managing invasions effectively 10,31.
Current control strategies have integrated mechanical, chemical, biological, and ecological approaches, with varying degrees of success depending on region-specific factors. Mechanical control, such as repeated mowing and tilling, is commonly used but often only provides temporary suppression unless followed by other treatments 1. Unlike in the United States, herbicidal control, particularly using glyphosate or imazapyr, is widely practiced and has shown effectiveness when applied persistently and in combination with burning or mowing 46. However, concerns over herbicide resistance and environmental impacts have heightened interest in integrated management approaches, in which prescribed burning can reduce biomass and enhance herbicide efficacy, although it may also stimulate regrowth if not carefully managed 46. Prescribed burning can reduce biomass and enhance herbicide efficacy but may also stimulate regrowth if not carefully managed. Biological control remains limited, though research into host-specific fungal pathogens and competitive native species shows promise 47. Restoration-based strategies that re-establish competitive native vegetation following control treatments are gaining traction, especially in conservation areas 46. Globally, successful long-term management requires adaptive, site-specific approaches that consider ecological, economic, and social dimensions.
This study provides ecoregional insights that can inform targeted management strategies for cogon grass. For instance, in previously cold-limiting ecosystems of northern Asia and Europe, early eradication efforts may prevent establishment. In agricultural landscapes of Africa and South America, sequential treatments combining mechanical removal and herbicide application could reduce spread and limit economic losses. In fire-prone savannas and grasslands of the southeastern United States, prescribed burning integrated with postfire herbicide treatments may help control post-disturbance expansion. By linking management strategies to specific regional contexts, these approaches can be calibrated to local ecological and climatic conditions, improving their effectiveness.
Limitations and future directions
This study mainly relied on bioclimatic and selected environmental variables (e.g., wildfire, land use, soil properties, HII), while excluding key ecological processes such as biotic interactions, competition with native species, and dispersal constraints. These omissions may reduce the accuracy of invasion risk predictions. In addition, our assessment is based on model projections without field validation, which limits confidence in the results. Future research should integrate long-term field data, mechanistic experiments, and remote sensing to validate and refine models. Incorporating ecological interactions and expanding field-based evidence will improve prediction accuracy and strengthen early detection and management strategies for cogon grass.
Conclusions
In this study, the global habitat suitability of cogon grass was assessed under current and future climate scenarios (SSP1-2.6 to SSP5-8.5) via SDMs based on five machine learning algorithms. Among these models, the MaxEnt model, in which environmental variables and wildfire data are incorporated, demonstrated the highest prediction accuracy. Currently, cogon grass has spread well beyond its native range, establishing itself in regions including Africa, southern Europe, South America, and the United States, with suitable habitats spanning between 40°N and 35°S. Under future climate change, its potential range is projected to expand significantly, particularly in Central Africa and South America, with a habitat suitability increase reaching 124.77% of current under the SSP5-8.5 scenario from 2061–2080. At present, extremely suitable habitats occur in 41 countries; this number is projected to increase to 70 countries. The results also highlight the critical role of climate change and wildfires in driving these changes, with 17 countries—including Sierra Leone, Rwanda, and Zaire—transitioning from unsuitable to highly suitable conditions. Currently, cogon grass is present in 64 distinct ecoregions and projections suggest that cogon grass may expand its habitat into up to 73 ecoregions in the future. This study offers insights into the future invasion risk of cogon grass and emphasizes the urgency of implementing effective prevention, legal regulation, and coordinated global efforts. Strengthening international and national collaboration is essential to safeguard ecosystems and economic assets against the accelerating threat of this invasive species.
Materials and methods
Global species occurrence data
A total of 29,159 occurrence records for cogon grass of all varieties were obtained from the Global Biodiversity Information Facility, GBIF)14, an open-access platform that provides species occurrence data compiled from a wide range of nonsystematic sources, including museum specimens and administrative datasets16. The GBIF plays a key role in promoting the sharing of biodiversity data for research, policy, and decision-making purposes for biodiversity conservation and sustainable management14. To increase model accuracy and minimize the effects of sampling bias and duplicate coordinates, we applied spatial rarefaction using SDM toolbox v.2.4 48,49 in ArcGIS 10.8 (Esri Korea, Seoul). We excluded records dated prior to 1970, data from unreliable sources, records lacking date information, and points located in ecologically irrelevant regions such as oceans, deserts, and polar zones. Expert consultation was also employed to validate the species occurrence records and finalize the species presence points of cogon grass for species distribution modeling. Rarefaction was performed at multiple spatial resolutions (1, 5, 10, 20, and 50 km) to eliminate duplicate records within each grid cell, which yielded six datasets, including one non-rarefied version. This process resulted in a reduced number of records: 13,780 records at 1 km, 5,934 records at 5 km, 3,637 at 10 km, 2,113 records at 20 km, and 1,006 records at 50 km (Fig. 1). These datasets were subsequently used for MaxEnt-based modeling of the cogon grass distribution.
Environmental variable selection
A
Bioclimatic variables such as temperature and precipitation are important for modeling the global spatial distribution of cogon grass (Table S1). Current climate data, representing the period from 1979 to 2013, were considered the baseline for current climatic conditions in the analysis process and were obtained from PaleoClim v1.250. Similarly, future climate projections were obtained from the WorldClim 2.1 database51. All the bioclimatic data exhibit a spatial resolution of 2.5 arc-minutes. We employed four shared socioeconomic pathway (SSP) scenarios, namely, SSP1-2.6, SSP2-4.5, SSP3-7.0, and SSP5-8.5, which represent a range of low to high greenhouse gas concentration trajectories. These scenarios were adopted to model potential climatic conditions for the period from 2061 to 2080. SSP describe potential trajectories of global societal development throughout the 21st century, thereby incorporating factors such as population growth, economic progress, technological advancement, and environmental challenges. Each pathway is based on a distinct socioeconomic narrative—ranging from sustainability-oriented development (SSP1) to fossil fuel–driven growth (SSP5) 52,53.
SSP1-2.6 represents a sustainability-focused pathway characterized by the rapid adoption of renewable energy and reduced dependence on fossil fuels. This scenario supports environmentally sustainable economic growth and is associated with a projected increase in the global temperature of approximately 2°C above preindustrial levels by 2100 42. SSP2-4.5 reflects an intermediate pathway, extending historical trends of uneven economic development and moderate climate policy implementation. Notably, partial achievement of emission reduction targets is assumed, resulting in projected warming of 2.7°C by 210052.
SSP3-7.0 is characterized by slow economic growth, limited technological advancement, continued heavy reliance on fossil fuels, and moderate population growth in developing countries. These conditions increase societal vulnerability to climate change impacts and are projected to yield a 3.6°C temperature increase by 210052,54. SSP5-8.5 entails the assumptions of high fossil fuel use, rapid technological innovation, and globalization, alongside energy-intensive economic growth and unregulated urban-centered development. Under this scenario, a temperature increases of 4.4°C is projected by 210042,52. These varied SSP scenarios, which represent a spectrum of moderate to extreme warming trajectories, offer robust frameworks for predicting potential changes in the distributions of invasive species. The bioclimatic data for the future scenarios were derived from the MPI-ESM1-2-HR global climate model 55 of the Coupled Model Intercomparison Project Phase 6 (CMIP6), which exhibits a consistent spatial resolution of 2.5 arcminutes (~ 4.5 km at the equator) 56.
Wildfires are critical components of terrestrial ecosystems and affect global carbon cycles, vegetation dynamics, and atmospheric composition while posing significant threats to biodiversity, human health, and socioeconomic stability 11. Wildfire can enhance the invasion of alien plants via the alteration in disturbance regimes, thereby increasing resources and competitive success over native plants33. To account for this factor, we downloaded wildfire data recorded between 2000 and 2024 by the Moderate Resolution Imaging Spectroradiometer (MODIS) from the Fire Information for Resource Management System (FIRMS) of the National Aeronautics and Space Administration 57 and Global Wildfire Information System 58.
Land use and land cover change are important in invasion ecology because they create suitable condition for promoting the introduction, colonization, and spread of invasive species by affecting niche availability and dispersal potential23,59. The land cover map used has seven categories including forest, agriculture land, grass land, wetland, non-vegetated land, mixed land, and artificial land (Food and Agriculture Organization database 60. The artificial land categories include human settlements, and infrastructure development.
Soil characteristics such as organic matter, carbon content, and pH play crucial roles in determining nutrient availability, microbial dynamics, and habitat suitability for invasive species38. Invasive species exhibit unique properties that allow them to modify soil characteristics61. These altered soils can provide a more favorable environment for the establishment of invasive species 62. 2016). Therefore, we incorporated three soil characteristics, i.e., soil pH, soil carbon, and soil moisture levels, into the model to better understand the distributions of cogon grass-related variables 60.
The human influence index (HII) encompasses various human activities, such as global trade and tourism, transportation and infrastructure development, urbanization and industrialization, and agriculture, that contribute to the disruption of natural biogeographical boundaries and promote the invasion of alien species45. These anthropogenic variables serve as essential ecological indicators for predicting species distributions under changing environmental conditions. Here, we incorporated HII data sourced from the Socioeconomic Data and Applications Center (SEDAC) of National Aeronautics and Space Administration 63.
We employed Pearson’s correlation analysis to reduce multicollinearity and increase model accuracy. We eliminated highly correlated environmental variables (Pearson’s r > 0.75 and p < 0.05). Finally, six bioclimatic variables, i.e., annual mean temperature (Bio1), mean diurnal range (Bio2), isothermality (Bio3), annual precipitation (Bio12), precipitation in the wettest month (Bio13), and precipitation in the driest month (Bio14), as well as five additional environmental variables, i.e., land cover change, wildfires, soil carbon, soil moisture, soil pH, and the HII, were selected for species distribution modeling of cogon grass (Table 1).
Species distribution modeling of cogon grass via machine learning algorithms
In this study, we employed five machine learning algorithms—boosted regression trees (BRTs), a generalized additive model (GAM), the maximum entropy (MaxEnt) model, a random forest (RF) model, and the extreme gradient boosting (XGBoost) model—to predict the global spatial distribution of cogon grass. Machine learning algorithms (MLAs) contribute significantly to species distribution modeling by facilitating the efficient processing of complex ecological datasets, thus increasing the accuracy of species occurrence predictions and promoting the resolution of complex ecological challenges64. These algorithms enable the analysis of the complex relationships between environmental variables and species occurrence, leading to a greater understanding of species distributions and supporting conservation and management efforts65.
The BRT model effectively resolves nonlinear relationships, identifies important predictor variables, accounts for interactions among predictors, and captures the relative importance of each predictor—tasks that are often challenging for traditional regression-based models66. The BRT model is applicable for modeling a wide range of organisms and environmental variables. GAMs constitute a semiparametric approach for predicting nonlinear relationships between environmental variables (e.g., temperature and chlorophyll) and species abundance, enabling reliable spatial predictions within specific environments67. The MaxEnt algorithm can be used to integrate species occurrence data with environmental variables, such as climate, land cover changes, and wildfires, to predict potential distributions in new regions on the basis of the principle of maximum entropy 68. It relies exclusively on species presence data, avoiding inaccuracies stemming from the use of unreliable absence data due to the unstable and expanding ranges of invasive species 69. The MaxEnt algorithm is widely applied in modeling both native and invasive species and is valuable in conservation and management decision-making processes19,59. The RF model is an effective ensemble classifier with several advantageous features. It accommodates diverse input variable types and generates numerous decision trees, determining classifications through majority voting 70. It effectively manages correlated variables and provides robust predictions. The XGBoost model, which leverages its advanced boosting framework, achieves high accuracy in feature selection and offers excellent scalability, making it particularly well suited for large-scale species distribution modeling applications71. We evaluated the model performance of each algorithm and selected the best algorithm for simulating cogon grass distributions under current and future climate change scenarios.
The data for alien and invasive species are often unreliable due to their expanding distribution ranges and the likelihood that they have not yet reached equilibrium, which can lead to misinterpretation in modeling and analysis69. Therefore, global background points (pseudoabsence) were generated using ArcGIS 10.8 (ESRI, Redlands, CA, USA), following methodologies outlined in previous studies72,73 (Barbet-Massin et al., 2012; Adhikari et al., 2018). BRT, GAM, MaxEnt, RF, and XGBoost modeling was performed by selecting a single model in the Biomod2 Package v.4.2–6.2 for GNU R (https://cran.r-project.org/web/packages/biomod2/index.html) 74. For model evaluation, the occurrence dataset was randomly split at a 3:1 ratio for model training and testing and 100 independent replications were performed to ensure the robustness of each species distribution model.
Model evaluation, validation and selection of the best modeling algorithm
To evaluate the prediction accuracy of each algorithm, five evaluation metrics, namely, the area under the receiver operating characteristic (ROC) curve (AUC) 75, true skill statistic (TSS)76, kappa, sensitivity, and specificity, were employed. The primary metric AUC is a threshold-independent indicator that represents the ability of a given model to differentiate between species presence and absence24. AUC values range from 0 to 1, with higher values indicating better model discrimination ability. On the basis of the classification proposed by Swets (1988), the AUC values were interpreted as follows: 0.5–0.6 (fail), 0.6–0.7 (poor), 0.7–0.8 (fair), 0.8–0.9 (good), and 0.9–1.0 (excellent) 24,77.
The TSS and kappa statistics are threshold-dependent metrics that capture both omission and commission errors, providing insight into the ability of the model to distinguish between species presence and absence 76. The values of both the TSS and kappa metrics range from − 1 to + 1 and account for both commission and omission errors. Values approaching + 1 indicate high prediction accuracy, whereas values near zero or negative values indicate poor model performance due to prediction errors 76.
Similarly, sensitivity captures the ability to correctly identify true presence, and specificity reflects the ability of the model to predict true absence. Despite being greatly affected by the choice of threshold—often resulting in inconsistent outcomes across presence-only datasets—sensitivity and specificity, when used together, provide a more reliable evaluation of model performance under varying threshold and prevalence conditions, aiding in the determination of potential biases and limitations 78. Notably, sensitivity and specificity values range from − 1 to + 1, indicating poor to perfect agreement. In this study, we employed these five-evaluation metrics and selected the best modeling algorithm for cogon grass. The selected algorithm was then applied in our analysis of the global spatial distribution and to assess habitat suitability in different countries worldwide.
Worldwide habitat suitability estimation for cogon grass
The binary distribution map, generated using the TSS thresholds defined during model calibration74, was employed to quantify suitable areas by country under baseline and projected climate change scenarios. The resulting model outputs indicate the global presence or absence of suitable area for cogon grass. ArcGIS Desktop 10.8 (Esri Korea, Seoul) was employed to map the habitat suitability of cogon grass across six continental regions and 195 countries. Similarly, changes from the historic distribution of suitable areas to the 2061–2080 period were analyzed for the four SSPs. The average habitat suitability of cogon grass was calculated for each country for current and projected future climate scenarios and classified into five categories, i.e., unsuitable (0), poorly suitable (0.01–0.25 of the country’s area), moderately suitable (0.25–0.5), highly suitable (0.5–0.75), and extremely suitable (0.75–1.0), across 195 countries worldwide.
Habitat suitability was also estimated across the seven land cover categories. Finally, we quantified the habitat coverage of cogon grass within global grassland areas and identified changes in the extent of invasible habitat in the African, Australian, and South American savannas. Our definition of grassland areas includes wide variety of grassland formation types, including alpine meadows, temperate steppes, tropical savannas, and semi-desert scrublands and was derived from the World Wildlife Fund 60.
Additional information
Conflict of interest
The authors have declared that no competing interests exist.
Ethical statement
No ethical statement was reported.
A
Funding
This work is supported by the Korea Environment Industry and Technology Institute (KEITI) through the Climate Change R&D Project for New Climate Regime, funded by the Korea Ministry of Environment (MOE) (RS-2022-KE002369, RS-2022-KE003570001) and Florida Fish and Wildlife Conservation Commission grant (22131).
A
Author Contribution
Pd.A. designed the research and methodology. Pd.A., A.P., Pb.A., E.S.L., D.R.B., and P.M.M. performed data collection, data curation, and analysis. Pd.A. and Y.H.L. carried out the modeling and validation of results. Pd.A. prepared the original manuscript. Pd.A. and J.T. reviewed and edited the manuscript. S.H.H. and C.S. supervised the study and managed the research funding. All authors read and approved the final manuscript.
Corresponding authors
Correspondence to Sun Hee Hong or Changwan Seo
Ethics declarations
Competing interests
The authors declare no competing interests.
Approval for human experiments
Not applicable
A
Data Availability
The datasets used in this study will be made available upon reasonable request to the corresponding authors.
Electronic Supplementary Material
Below is the link to the electronic supplementary material
References
1.
MacDonald, G. E. Cogongrass (Imperata cylindrica)—biology, ecology, and management. Crit. Rev. Plant. Sci. 23, 367–380 (2004).
2.
Kato-Noguchi, H. Allelopathy and allelochemicals of Imperata cylindrica as an invasive plant species. Plants 11, 2551 (2022).
3.
Parker, C. Parasitic weeds: a world challenge. Weed Sci. 60, 269–276 (2012).
4.
Hubbard, C. Imperata cylindrica taxonomy, distribution, economic significance and control. Ch. 1, 5–13Imperial Forestry Bureau, Oxford, Great Britain, (1944).
5.
GISD. Vol. (IUCN/SSC Invasive Species Specialist Group, 2024). (2024).
6.
Lucardi, R. D., Wallace, L. E. & Ervin, G. N. Patterns of genetic diversity in highly invasive species: Cogongrass (Imperata cylindrica) expansion in the invaded range of the southern United States (US). Plants 9, 423 (2020).
7.
Lippincott, C. L. Effects of Imperata cylindrica (L.) Beauv. (cogongrass) invasion on fire regime in Florida sandhill (USA). Nat. Areas J. 20, 140–149 (2000).
8.
Holzmueller, E. J. & Jose, S. Response of the invasive grass Imperata cylindrica to disturbance in the southeastern forests, USA. Forests 3, 853–863 (2012).
9.
Estrada, J. A. & Flory, S. L. Cogongrass (Imperata cylindrica) invasions in the US: mechanisms, impacts, and threats to biodiversity. Glob Ecol. Conserv. 3, 1–10 (2015).
10.
Parker, C. Imperata cylindrica (cogon grass). CABI Compendium. 10.1079/cabicompendium.28580 (2022).
11.
Wasserman, T. N. & Mueller, S. E. Climate influences on future fire severity: a synthesis of climate–fire interactions and impacts on fire regimes, high-severity fire, and forests in the western United States. Fire Ecol. 19, 1–22 (2023).
12.
North, M. P. et al. Strategic fire zones are essential to wildfire risk reduction in the western United States. Fire Ecol. 20, 50 (2024).
13.
Zhou, J., Hu, J., Liu, J. & Zhang, W. Elucidating the gastroprotective mechanisms of Imperata cylindrica Beauv. var. major (Nees) C. E. Hubb through UHPLC-MS/MS and systems network pharmacology. Sci. Rep. 14, 27815 (2024).
14.
GBIF. (Global Biodiversity Information Facility, Copenhagen & Denmark (2024). https://www.gbif.org/species/8424006
15.
Divate, N., Solís, D., Thomas, M. H., Alvarez, S. & Harding, D. An economic analysis of the impact of cogongrass among nonindustrial private forest landowners in Florida. Sci. 63, 201–208 (2016).
16.
Adhikari, P. et al. Global assessment of invasion risk: Ardisia elliptica, one of the most noxious tropical shrubs in the world. Ecol. Process. 14, 55 (2025).
17.
Elith, J. & Leathwick, J. R. Species distribution models: ecological explanation and prediction across space and time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697 (2009).
18.
Adhikari, P., Kim, B. J., Hong, S. H. & Lee, D. H. Climate change induced habitat expansion of nutria (Myocastor coypus) in South Korea. Sci. Rep. 12, 1–12 (2022).
19.
McCulloch-Jones, E. J., Kraaij, T., Crouch, N. & Faulkner, K. T. Assessing the invasion risk of traded alien ferns using species distribution models. NeoBiota 87, 161–189 (2023).
20.
Adhikari, P. et al. Global invasion risk assessment of Lantana camara, a highly invasive weed, under future environmental change. Glob Ecol. Conserv. 55, e03212 (2024).
21.
Poudel, A. et al. Assessing the potential distribution of Oxalis latifolia, a rapidly spreading weed, in East Asia under global climate change. Plants 12, 3254 (2023).
22.
Hong, S. H., Lee, Y. H., Lee, G., Lee, D. H. & Adhikari, P. Predicting impacts of climate change on northward range expansion of invasive weeds in South Korea. Plants 10, 1604 (2021).
23.
Williams, D. A. et al. Predictor importance in habitat suitability models for invasive terrestrial plants. Divers. Distrib. 30, e13906 (2024).
24.
Thuiller, W., Lavorel, S. & Araújo, M. B. Niche properties and geographical extent as predictors of species sensitivity to climate change. Glob Ecol. Biogeogr. 14, 347–357. 10.1111/j.1466-822X.2005.00162.x (2005).
25.
Adhikari, P. et al. Potential impact of climate change on plant invasion in the Republic of Korea. J. Ecol. Environ. 43, 36. 10.1186/s41610-019-0134-3 (2019).
26.
Gajendiran, K., Kandasamy, S. & Narayanan, M. Influences of wildfire on the forest ecosystem and climate change: A comprehensive study. Environ. Res. 240, 117537 (2024).
27.
Aslan, C. E. & Dickson, B. G. Non-native plants exert strong but under-studied influence on fire dynamics. NeoBiota 61, 47–64 (2020).
28.
Leal-Medina, C., Lopatin, J., Contreras, A., González, M. & Galleguillos, M. Post-fire Pinus radiata invasion in a threatened biodiversity hotspot forest: a multi-scale remote sensing assessment. Ecol. Manag. 561, 121861 (2024).
29.
Butler, O. M., Lewis, T. & Chen, C. Do soil chemical changes contribute to the dominance of blady grass (Imperata cylindrica) in surface fire-affected forests? Fire 4, 23 (2021).
30.
Box, R. C. International Handbook of Disaster Research Vol. 1–16 (Springer, 2023).
31.
USDA. After the Fire. Forest Service & United States Department of Agriculture., (2025). https://afterthefireusa.org/ Accessed April 16, 2025.
32.
Reilly, M. J. et al. Repeated, high-severity wildfire catalyzes invasion of non-native plant species in forests of the Klamath Mountains, northern California, USA. Biol. Invasions. 22, 1821–1828 (2020).
33.
Wimberly, M. C., Wanyama, D., Doughty, R., Peiro, H. & Crowell, S. Increasing fire activity in African tropical forests is associated with land use and climate change. Authorea Preprints (2023).
34.
Kraaij, T., Baard, J. A., Arndt, J., Vhengani, L. & Van Wilgen, B. W. An assessment of climate, weather, and fuel factors influencing a large, destructive wildfire in the Knysna region, South Africa. Fire Ecol. 14, 1–12 (2018).
35.
Andela, N. et al. A human-driven decline in global burned area. Science 356, 1356–1362 (2017).
36.
Giorgis, M. A. et al. A review of fire effects across South American ecosystems: the role of climate and time since fire. Fire Ecol. 17, 1–20 (2021).
37.
Becklin, K. M. et al. Examining plant physiological responses to climate change through an evolutionary lens. Plant. Physiol. 172, 635–649 (2016).
38.
Nievas, R. P., Calderon, M. R. & Moglia, M. M. Environmental factors affecting the success of exotic plant invasion in a wildland–urban ecotone in temperate South America. Neotrop. Biol. Conserv. 14, 257–274 (2019).
39.
Finch, D. M. et al. Effects of climate change on invasive species. In Invasive Species in Forests and Rangelands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector 57–83 (2021).
40.
Chen, P. et al. Deterministic responses of biodiversity to climate change through exotic species invasions. Nat. Plants. 10, 1464–1472 (2024).
41.
Oh, M., Heo, Y., Lee, E. J. & Lee, H. Major environmental factors and traits of invasive alien plants determining their spatial distribution. J. Ecol. Environ. 45, 1–10 (2021).
42.
Lee, J. Y. et al. Future global climate: scenario-based projections and near-term information. (2021).
43.
Rasool, G. et al. Projecting climate change impact on precipitation patterns during different growth stages of rainfed wheat crop in the Pothwar Plateau, Pakistan. Climate 12, (2024).
44.
Rad, S. P. H. et al. Predicting the spread of invasive Imperata cylindrica under climate change: a global risk assessment and future distribution scenarios. PLoS One. 20, e0321027 (2025).
45.
Gallardo, B., Zieritz, A. & Aldridge, D. C. The importance of the human footprint in shaping the global distribution of terrestrial, freshwater, and marine invaders. PLoS One. 10, e0125801 (2015).
46.
Lebrun, A. Cogongrass Control Program in the Southeastern United States – Alabama, Georgia, Mississippi and South Carolina (US Department of Agriculture, Riverdale, 2020).
47.
Purnama, H. et al. Potential biological control agents for management of cogongrass (Cyperales: Poaceae) in the southeastern USA. (2016).
A
48.
Brown, J. L., Bennett, J. R. & French, C. M. SDMtoolbox 2.0: the next-generation Python-based GIS toolkit for landscape genetic, biogeographic and species distribution model analyses. PeerJ 5, e4095. 10.7717/peerj.4095 (2017).
A
49.
Boria, R. A., Olson, L. E., Goodman, S. M. & Anderson, R. P. Spatial filtering to reduce sampling bias can improve the performance of ecological niche models. Ecol. Model. 275, 73–77. 10.1016/j.ecolmodel.2013.12.012 (2014).
50.
Brown, J. L., Hill, D. J., Dolan, A. M., Carnaval, A. C. & Haywood, A. M. PaleoClim: high spatial resolution paleoclimate surfaces for global land areas. Sci. Data. 5, 1–9 (2018).
51.
Fick, S. E. & Hijmans, R. J. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302–4315 (2017).
52.
Riahi, K. et al. The shared socioeconomic pathways and their energy, land use, and greenhouse gas emissions implications: an overview. Glob Environ. Change. 42, 153–168 (2017).
53.
O’Neill, B. C. et al. The Scenario Model Intercomparison Project (ScenarioMIP) for CMIP6. Geosci. Model. Dev. 9, 3461–3482 (2016).
A
54.
O’Neill, B. C. et al. The roads ahead: narratives for shared socioeconomic pathways describing world futures in the 21st century. Glob Environ. Change. 42, 169–180 (2017).
55.
Gutjahr, O. et al. Max Planck Institute Earth System Model (MPI-ESM1.2) for the High-Resolution Model Intercomparison Project (HighResMIP). Geosci. Model. Dev. 12, 3241–3281 (2019).
56.
Eyring, V. et al. Overview of the Coupled Model Intercomparison Project Phase 6 (CMIP6) experimental design and organization. Geosci. Model. Dev. 9, 1937–1958 (2016).
57.
NASA. Fire Information for Resource Management System. (2025). https://firms.modaps.eosdis.nasa.gov/
58.
GWIS. National Aeronautics and Space Administration and European Commission, (2025). https://gwis.jrc.ec.europa.eu
59.
Adhikari, P., Kim, H. W., Shin, M. S., Hong, S. H. & Cho, Y. Potential distribution of the silver stripped skipper (Leptalina unicolor) and maiden silvergrass (Miscanthus sinensis) under climate change in South Korea. Entomol Res (2022).
60.
FAO. (Food and & Organization, A. (2025). https://www.fao.org/ (Accessed 23 February 2025).
61.
Szumańska, I. et al. Invasive plant species distribution is structured by soil and habitat type in the city landscape. Plants 10, 773 (2021).
62.
Kuebbing, S. E. & Nuñez, M. A. Invasive non-native plants have a greater effect on neighbouring natives than other non-natives. Nat. Plants. 2, 1–7 (2016).
63.
SEDAC. (National Aeronautics and Space Administration (NASA). (2025).
64.
Beery, S., Cole, E., Parker, J., Perona, P. & Winner, K. In Proc. 4th ACM SIGCAS Conf. Computing and Sustainable Societies, 329–348.
65.
Refaat, A. M., Youssef, A. M., Mosallam, H. A. A. & Farouk, H. Predicting the effect of climate change on the spatiotemporal distribution of two endangered plant species, Silene leucophylla Boiss. and Silene schimperiana Boiss., using machine learning in Saint Catherine Protected Area, Egypt. Beni-Suef Univ. J. Basic. Appl. Sci. 13, 98 (2024).
66.
Yu, H., Cooper, A. R. & Infante, D. M. Improving species distribution model predictive accuracy using species abundance: Application with boosted regression trees. Ecol. Model. 432, 109202 (2020).
67.
Drexler, M. & Ainsworth, C. H. Generalized additive models used to predict species abundance in the Gulf of Mexico: an ecosystem modeling tool. PLoS One. 8, e64458 (2013).
68.
Phillips, S. J., Anderson, R. P. & Schapire, R. E. Maximum entropy modeling of species geographic distributions. Ecol. Model. 190, 231–259 (2006).
69.
Jiménez-Valverde, A. et al. Use of niche models in invasive species risk assessments. Biol. Invasions. 13, 2785–2797. 10.1007/s10530-011-9963-4 (2011).
70.
Breiman, L. Random forests. Mach. Learn. 45, 5–32 (2001).
71.
Effrosynidis, D., Tsikliras, A., Arampatzis, A. & Sylaios, G. Species distribution modelling via feature engineering and machine learning for pelagic fishes in the Mediterranean Sea. Appl. Sci. 10, 8900 (2020).
72.
Barbet-Massin, M., Jiguet, F., Albert, C. H. & Thuiller, W. Selecting pseudo-absences for species distribution models: how, where and how many? Methods Ecol. Evol. 3, 327–338 (2012).
73.
Adhikari, P. et al. Potential impact of climate change on the species richness of subalpine plant species in the mountain national parks of South Korea. J. Ecol. Environ. 42, 36 (2018).
74.
Thuiller, W. et al. biomod2: Ensemble platform for species distribution modeling. Version 4.2-6-2 (2025).
75.
Pearson, R. G. Species’ distribution modeling for conservation educators and practitioners. Lessons Conserv. 3, 54–89 (2010).
76.
Allouche, O., Tsoar, A. & Kadmon, R. Assessing the accuracy of species distribution models: prevalence, kappa and the true skill statistic (TSS). J. Appl. Ecol. 43, 1223–1232 (2006).
77.
Swets, J. A. Measuring the accuracy of diagnostic systems. Science 240, 1285–1293. 10.1126/science.3287615 (1988).
78.
Leroy, B. et al. Without quality presence–absence data, discrimination metrics such as TSS can be misleading measures of model performance. J. Biogeogr. 45, 1994–2002 (2018).
Figure 1.
Figure 2.
Figure 3.
Figure 4.
Figure 5.
Total words in MS: 9529
Total words in Title: 19
Total words in Abstract: 193
Total Keyword count: 7
Total Images in MS: 5
Total Tables in MS: 6
Total Reference count: 78