Environmental Science and Pollution Research (2022) 29:63260–63276

https://doi.org/10.1007/s11356-022-20054-y

RESEARCH ARTICLE

Habitat change and biodiversity loss in South and Southeast Asian

countries
Yan‑Ling Tan1 · Jen‑Eem Chen2 · Thian‑Hee Yiew3 · Muzafar Shah Habibullah4

Received: 28 October 2021 / Accepted: 29 March 2022 / Published online: 22 April 2022
© The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2022

Abstract
South and Southeast Asia is by far the most populous region in Asia, with the greatest number of threatened species. Changes
in habitat are a major contributor to biodiversity loss and are more common as a result of land-use changes. As a result, the
goal of this study is to use negative binomial regression models to investigate habitat change as one of the important drivers
of biodiversity loss in South and Southeast Asian countries from 2013 to 2018. According to the negative binomial estimates,
the findings for the habitat change measures are quantitatively similar for the impacts of agricultural land and arable land on
biodiversity threats. Agricultural and arable land both have a positive impact on biodiversity loss. We found that, contrary to
our expectations, the forest area appears to have an unexpected direct influence on the number of threatened species. A higher
number of threatened species is associated with rising per capita income, human population and a low level of corruption
control. Finally, the empirical findings are consistent across taxonomic groups, habitat change measures and Poisson-based
specifications. Some policy implications that could mitigate biodiversity loss include educating and promoting good govern-
ance among the population and increase the conservation effort to sustain green area and national forest parks in each country.

Keywords Biodiversity loss · Habitat change · Land-use changes

Introduction put biodiversity in jeopardy due to rapid urbanisation and

resource exploitation. Biodiversity loss has emerged as the
Biodiversity is essential for providing genetic resources for most serious global environmental challenge confronting
all livestock, plants and marine species harvested for food the world today. As a result, worldwide concerns about
(Kaur et al. 2019). Nonetheless, economic activities have the rapid extinction of species have been heightened (Li
et al. 2013). Global species extinction rates are expected to
increase 1000 times faster than the background rate (Pimm
Responsible Editor: Eyup Dogan
et al. 2014).
* Jen‑Eem Chen In recent years, developing countries in Asia have demon-
jechen@uitm.edu.my strated resilient economic growth, but this has also exposed
Yan‑Ling Tan them to the greatest threats to their biodiversity. Figure 1
tanya163@uitm.edu.my depicts the decline in biodiversity in Asia during 2013 and
Thian‑Hee Yiew 2020. It is clear to observe that the number of threatened
yiewth@utar.edu.my species was increasing in each region of Asia. Additionally,
Muzafar Shah Habibullah South and Southeast Asia, which have the majority of their
muzafar@putrabs.edu.my countries covered in tropical rainforests that provide habitat
1 for species, experienced the highest number of threatened
Faculty of Business and Management, Universiti Teknologi
MARA Cawangan Johor, Kampus Segamat, Malaysia species and also a sharp increase in the biodiversity loss. The
2 International Union for the Conservation of Nature classifies
Faculty of Business and Management, Universiti Teknologi
MARA Cawangan Perlis, Kampus Arau, Malaysia several local species as vulnerable (VU), endangered (EN)
3 or critically endangered (IUCN 2010).
Faculty of Business and Finance, Universiti Tunku Abdul
Rahman, Kampar, Malaysia Furthermore, as shown in Fig. 2, Malaysia experienced
4 the greatest loss of species, followed by Indonesia and
Putra Business School, Selangor, Malaysia

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Environmental Science and Pollution Research (2022) 29:63260–63276 63261

India, where over 1000 species were threatened, while
Timor-Leste had the least threatened species. Furthermore,
a large number of species have become extinct in three
mega diverse countries, Malaysia, Indonesia and the Phil-
ippines, which house the majority of the world’s flora and
fauna. Malaysia, Indonesia and India have a total of 1351,
1348 and 1049 endangered species, respectively.
Alternatively, Fig. 3 depicts the number of threatened
species in South and Southeast Asia by taxonomic group for
the years 2013 and 2020. In 2013, the number of threatened

Fig. 1  Number of threatened

species in Asian regions, 2013 Number of threatened species 2013 2020
and 2020. Source: IUCN (2020, 10,000 9,459
version 1) and IUCN (2013,
version 2) 8,000 7,456

6,000

4,000 3,010
2,380
2,024 1,861
2,000
264 474
0
South & Southeast Asia West & Central Asia East Asia North Asia

Fig. 2  Average number of

threatened species in South Timor-Leste 27
and Southeast Asian countries, Bhutan 80
2013–2020. Source: IUCN Nepal 104
(various versions) Bangladesh 148
Brunei 212
Laos 219
Cambodia 255
Singapore 300
Myanmar 317
Sri Lanka 598
Thailand 617
Vietnam 627
Philippines 786
India 1,049
Indonesia 1,348
Malaysia 1,351
0 100 200 300 400 500 600 700 800 900 1,000 1,100 1,200 1,300 1,400
Average number of threatened species

Fig. 3  Number of threatened

species by taxonomic groups Number of threatened species
10,000
in South and Southeast Asian 9,274
2013 2020
countries, 2013 and 2020. 9,000
Source: IUCN (various ver-
8,000
sions) “For reptiles, fishes,
molluscs, other invertebrates 7,000 7,304
and plants: the figure displayed 6,000
should be interpreted as the
number of species known to be 5,000

threatened within those species 4,000 3,555

that have been assessed to date,
3,000
and not as the overall total num-
1,863 2,557
ber of threatened species for 2,000 1,308
each group (IUCN 2020, p. 1)” 1,000
823 817 441 297 1,651
159
799 646 343 294 914 100
0
Mammals Birds Reptiles Amphibians Fishes Molluscs Other inverts Plants Total

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63262 Environmental Science and Pollution Research (2022) 29:63260–63276

species in South and Southeast Asia ranged from 100 for
molluscs to 2557 for plants. In 2020, a total of 9274 species
were classified as threatened. Plants, other invertebrates and
fishes suffered the greatest loss of species among all threat-
ened species. As a result, natural habitats in the tropical
rainforest were disappearing at an alarming rate.
Figure 4 depicts the trends in agricultural land, arable
land and forest area in countries in South and Southeast
Asia. As shown in Fig. 4(a), the total agricultural land was
327 million hectares in 2013, while the total arable land
was 237 million hectares. In 2018, agricultural and arable
land increased to 335 million hectares and 241 million
hectares, respectively. At the same time, Fig. 4(b) presents
the agricultural land, arable land and forest area by coun-
tries in 2018. India alone possessed 180 million hectares
of agricultural land and 156 million hectares of arable
land. Indonesia has 62 million hectares of agricultural
land and 26 million hectares of arable land, and Thailand
comes next with 22 million hectares of agricultural land
and 17 million hectares of arable land. Aside from that,
forest area in South and Southeast Asian countries is likely
to have decreased from 298 million hectares in 2013 to
294 million hectares in 2018, representing a 4.3 million
hectares loss (Fig. 4(a)).
This study differs from previous studies in three
aspects. First, empirical evidence about the effects of
habitat change in a larger context as a result of land-
use changes is presented. Changes in habitat can lead
to an increase in biodiversity loss in three ways: (i)
agricultural land and arable land changes such as
vegetation clearing for agricultural and other devel-
opmental purposes (Asafu-Adjaye 2003; MEA 2005;
(ii) reduction in forest cover, due to population and
employment pressures on forest cover, has resulted in
high fuelwood use, house building and land clearing
(Xu et al. 2006). Forest cover in protected areas, on
the other hand, reflects an initiative to protect biodi-
versity and human welfare; and (iii) ineffective forest
management in managing and optimising resources (Xu
et al. 2006). This has resulted in more physical places
to meet human demands, resulting in increased species
habitat loss and biodiversity loss (Asafu-Adjaye 2003;

Fig. 4  Agricultural land, arable

land and forest area in South '000 hectares
400,000.0
and Southeast Asian countries. 2013 2018
335,155.8
Source: WDI (2021) (a) Agri- 350,000.0
cultural land, arable land, and 293,914.2
300,000.0
forest area, 2013 and 2018. (b) 240,983.8
250,000.0
Agricultural land, arable land,
and forest area by countries in 200,000.0
2018 150,000.0
327,420.8
298,193.6
236,981.3
100,000.0

50,000.0

0.0
Agricultural land Arable land Forest area

(a)

'000 hectares
180,000
170,000
160,000 Agricultural land Arable land Forest area
150,000
140,000
130,000
120,000
110,000
100,000
90,000
80,000
70,000
60,000
50,000
40,000
30,000
20,000
10,000
0

(b)

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Environmental Science and Pollution Research (2022) 29:63260–63276
Hu et al. 2021). The indicator of land-use change is
frequently used to identify biodiversity threats (Hu
et al. 2021).
There are several aspects of land use such as agricultural
land, arable land and forest area (FAO 2021; WDI 2021).
Extensive land use may result in soil nutrient loss, land
degradation, grassland destruction and deforestation. Sug-
arcane farming practices, for example, have been found to
reduce soil organic matter and contribute to soil degrada-
tion in South Africa (Bailey et al. 2016). For simplicity,
agricultural land is the land area dedicated to permanent
crops, whereas arable land is dedicated to temporary crops
(FAO 2021; WDI 2021). However, our understanding of the
impact of habitat change is still limited because previous
studies did not distinguish between land-use conversion and
habitat change (for instance, permanent or temporary crops).
As a result, this study distinguished three indicators of habi-
tat change: agricultural land, arable land and forest area.
Secondly, previous studies used only threatened fish
species (Clausen and York 2008), only threatened plant
species (Habibullah et al. 2016), only threatened bird spe-
cies (Habibullah et al. 2018) or threatened species, species
richness and endemic species for mammals and birds (Kerr
and Burkey 2002; Kerr and Currie 1995; McPherson and
Nieswiadomy 2005) to capture the disaggregated effects of
biodiversity loss. We argued that habitat change and other
anthropogenic factors could cause individual species to
become endangered. Following that, regression analyses
were carried out on both the aggregated threatened species
and the individual threatened species, using method simi-
lar to those used in previous study (Naidoo and Adamow-
icz 2001). Individual threatened species data contains more
informative and complete information than threatened spe-
cies analysis using aggregated data.
Finally, we used annual panel data for 16 South and
Southeast Asian countries from 2013 to 2018 rather than
cross-sectional data, which has been used in several empiri-
cal studies (Asafu-Adjaye 2003; Kerr and Currie 1995;
Naidoo and Adamowicz 2001; Habibullah et al. 2021), or
data averaging (Clausen and York 2008). This is due to an
increase in the number of threatened species in South and
Southeast Asian countries. As a result of these shortcom-
ings, we decided to investigate habitat change as one of the
major drivers of biodiversity loss in South and Southeast
Asian countries from 2013 to 2018.
Literature review
Biological diversity (biodiversity) is defined as variation
among living organisms from all sources, including ter-
restrial, marine and other aquatic ecosystems, as well as
the ecological complexes that they form (MEA 2005).
63263
Biodiversity is essential for the proper functioning of eco-
systems, which support the delivery of ecosystem services,
that provide us with a variety of direct and indirect social
and economic benefits (Halkos 2011). Climate regulation,
food and water supply, disease and natural disaster control,
recreation, soil formation and nutrient cycling are among the
benefits (MEA 2005). Nonetheless, human activities have
consistently resulted in biodiversity loss and ecosystem dis-
ruption, with species becoming extinct at an unprecedented
rate. Eventually, biodiversity loss endangers human well-
being and, possibly, human survival (Cardinale et al. 2012;
Naeem et al. 2016; Skogen et al. 2018).
Biodiversity loss occurs when a specific diversity declines
due to species extinction or when the potential of the com-
ponents of diversity to provide a specific service is reduced
due to unsustainable harvesting (MEA 2005). Human devel-
opment and biodiversity loss are widely regarded as today’s
two major environmental challenges. Land conversion to
agriculture, air and water pollution, overexploitation, climate
change and urban development are all factors contributing
to a decrease in natural habitat area. Furthermore, the rate
of conversion of land to agriculture is much higher in the
tropical region (Perrings and Halkos 2015). This obviously
results in habitat loss caused by crop and pasture lands being
relocated (Deininger and Minten 2002; Tilman 1999; Tilman
et al. 1994).
One of the signs of environmental degradation that the
world is facing today is the loss of biodiversity. Biodiversity
loss involves the irreversible loss of valuable ecosystems,
and biodiversity decline is classified as a special issue in
environmental degradation (Asafu-Adjaye 2003). Rapid
urbanisation has increased the demand for natural resources,
resulting in the destruction of forests and natural habitat.
A large number of species are expected to decline or even
become extinct as a result of habitat loss by 2050 (Kong
et al. 2021; Powers and Jetz 2019). As a result, habitat loss
is a primary cause of biodiversity decline, and it is most
closely related to land-cover change. Conversions of land
for agricultural and other uses occur because of increasing
population growth, which has put the ecosystem in jeop-
ardy. According to Perrings and Halkos (2015), most of the
population in developing countries still lives in rural areas
and involves in agricultural activities that exploit natural
resources.
Agricultural land conversion or cultivated systems cover
roughly one-quarter of the Earth’s land surface (MEA 2005).
Between now and 2050, it is expected that 10–20% of exist-
ing grassland and forestland will be used for other purposes,
primarily agricultural land expansion as well as city and
infrastructure expansion (MEA 2005). Furthermore, due to
patchy conversion patterns, existing forest fragmentation
occurs, resulting in a reduction in the number and abun-
dance of species that can be supported on unconverted land
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63264
(Laurance et al. 2011; Lewis et al. 2009). As a result, habitat
destruction, degradation and fragmentation because of land-
use change are more important drivers of biodiversity loss
(Maxwell et al. 2016; Skogen et al. 2018). Extensive studies
have been conducted in recent years to investigate biodiver-
sity loss from a scientific standpoint. Nonetheless, quantita-
tive research in social science and economic contexts is still
constrained, owing primarily to data availability.
The previous studies investigated biodiversity loss in
various ways, including data, methodology and interpreta-
tions. For example, a few indicators were used to represent
biodiversity loss which were deforestation, the National
Biodiversity Risk Assessment Index (NABRAI) and the
number of threatened species. Deforestation was attempted
by Cropper and Griffiths (1994). On the other hand, the
National Biodiversity Risk Assessment Index (NABRAI)
was developed by Reyers et al. (1998), which takes into
account indices of pressure, state and response. Last but not
the least, the threatened species listed on the IUCN’s red list
were used to assess biodiversity loss (Gren et al. 2015; Kerr
and Currie 1995; Naidoo and Adamowicz 2001; Habibullah
et al. 2019; Habibullah et al. 2021). Human development
has directly resulted in habitat disturbances due to changes
in land use. Habitat disturbances include habitat loss due to
reduction in area and habitat fragmentation, which occurs
when a large habitat is divided into small parts (Kong
et al. 2021).
The consequences of habitat loss include population loss
and a reduction in species richness (Butchart et al. 2010;
Pimm et al. 2014). A large number of species are expected
to decline or even become extinct as a result of habitat loss
by 2050 (Powers and Jetz 2019). As a result, there is an
urgent need to examine biodiversity loss from the stand-
point of land use. In fact, land can be classified into two
types: human-dominated land and natural-dominated land
(Kong et al. 2021). Human-dominated land is used for
intensive human activities such as agriculture and urbanisa-
tion, whereas nature-dominated land, such as forests, grass-
land, water bodies and bare areas, are not directly altered
by humans. The effects of conversion of nature-dominated
land to urban land on biodiversity have been extensively
discussed (Güneralp and Seto 2013; He et al. 2014; Liu
et al. 2019).
However, until recently, the effects of agricultural expan-
sion on habitat loss were not given enough attention in bio-
diversity hotspots (Kehoe et al. 2015; Molotoks et al. 2018).
To some extent, it is argued that agricultural land provides
important habitat for terrestrial species (Foley et al. 2011).
Despite this, agrochemical use and agricultural mechanisa-
tion (Davis et al. 2017; Zermeño-Hernández et al. 2015),
seasonality in agricultural production and uncertainty in
agricultural structural adjustments are detrimental to bio-
logical survival and reproduction (Kong et al. 2021).
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Environmental Science and Pollution Research (2022) 29:63260–63276
Numerous studies have been conducted to investigate the
effects of land-use change and socioeconomic factors on bio-
diversity loss. Cropper and Griffiths (1994) used deforesta-
tion as a proxy for biodiversity in their early studies and used
fixed effects estimation to examine the effects of population
pressure on biodiversity in Africa, Latin America and Asia.
Based on the findings, rural population density increases
deforestation in Africa, whereas log price has a significant
impact on deforestation in Latin America. Furthermore, in
Africa and Latin America, there is a hump-shaped relation-
ship between per capita income and deforestation. However,
the independent variables are insignificant in explaining
deforestation in Asia, which may be due to the importance
of forest plantations in this continent.
Kerr and Currie (1995) were, to the best of our knowl-
edge, the first paper to use threatened species to assess
biodiversity loss. In 90 countries, they assessed the effects
of human activity on global extinction risk. The findings
revealed that human population poses a significant threat
to bird species, whereas per capita income poses a threat
to mammal species. Similar finding on the former was also
found by Habibullah et al. (2016) indicating that increase
in population growth increases the number of threatened
plant species in 143 countries. Protected areas, on the other
hand, have a strong correlation with mammalian population
density. More importantly, there is no evidence to support
the claim that species extinction is caused by habitat loss in
areas where the land-use pattern is statistically insignificant.
Kerr and Burkey (2002) then conducted another study,
this time focusing on the patterns of endemism and the
numbers of threatened birds and mammals in 42 countries
with tropical moist forests. They reported that mammals
are more threatened than birds in these countries, implying
that mammals are especially vulnerable to environmental
changes. In contrast to their previous study, Kerr and Currie
(1995) asserted that protected areas do not correspond to
threatened species because reserves are too small to provide
adequate protection for vertebrates, and reserve networks
are not related to threatened species at a large spatial scale.
Furthermore, human population growth rates and rising debt
levels in these countries should be prioritized for improve-
ment by developed countries.
Given that the data on threatened species has recently
become more readily available, an increasing number of
studies are using them as a biodiversity loss indicator. Fur-
thermore, the importance of land-use factors is gaining
attraction among academics, as habitat change is claimed to
be a primary driver of biodiversity loss. Naidoo and Adamo-
wicz (2001) have expanded the list of threatened species by
including plants, mammals, amphibians, reptiles, fishes and
invertebrates, as well as land-use factors. Using count-data
regression, they examined the relationship between threat-
ened species and per capita income in 100 countries. In five
Environmental Science and Pollution Research (2022) 29:63260–63276
of the seven taxonomic groups studied, the number of threat-
ened species was found to be related to per capita income.
In addition, land-use factors have a significant impact on
threatened species. Protected areas are important in prevent-
ing the extinction of amphibian species, whereas agricultural
land is a threat to invertebrate species.
Asafu-Adjaye (2003) found that protected land areas
could save species from extinction, particularly mammal
species, bird species and plant species, whereas land devel-
oped for agriculture and other uses only affects mammal
species. Aside from that, economic growth, population
density, number of tourists, good governance, exchange
rate and economic freedom all play a role in biodiversity
loss. According to Li et al. (2013), more arable land, higher
population density and economic growth will exacerbate
species loss in the long run. Perrings and Halkos (2015)
investigated the impact of agricultural productivity growth
and agricultural land conversion on threats to mammalian,
bird and plant species in 27 Sub-Saharan African countries.
They discovered that widespread agricultural expansion
is associated with increasing threats to biodiversity at all
time scales. On long time scales, agricultural intensifica-
tion is associated with a significant reduction in the threat
to all species.
Despite the fact that economic growth is commonly asso-
ciated with biodiversity declines, Dietz and Adger (2003)
and Gren et al. (2015) found no evidence to support this
claim. Nonetheless, Dietz and Adger (2003) proposed that
protected land and regulations governing the trade in endan-
gered species are effective biodiversity conservation initia-
tives. With respect to economic growth, on the contrary,
Habibullah et al. (2016, 2018) found that economic growth
showed negative impact on biodiversity loss in a sample of
over 140 countries. Furthermore, they also found that the
number of tourist arrivals impacted positively on the number
of threatened species in these countries.
Similarly, before further increases in economic pros-
perity result in reductions in species threat, a certain level
of institutional capacity (corruption and democracy) is
required (Gren et al. 2015). As a matter of fact, the work
by Habibullah et al. (2016, 2019) indicated that countries
that practices good governance experienced decreasing
level of biodiversity loss. Meanwhile, income inequality
would result in an increase in the number of threatened
species (Halkos 2011; Halkos and Tzeremes 2010; Mik-
kelson et al. 2007).
Although there are empirical studies that show the
land-use change could lead to biodiversity loss, some
studies show conflicting results. Pandit and Laband
(2007), for example, claimed that the relationship
between species richness and protected land area is
not well established across 127 countries. Despite
this, empirical results show that human population
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concentration is a critical factor affecting amphibians,
reptiles and bird species. Gren et al. (2015) concluded
that the effects of protected areas on biodiversity loss
are mixed. Protected areas may reduce the number of
threatened mammals, amphibians and reptiles while
increasing the number of threatened plant species. These
could be the results of ineffective management (Watson
et al. 2014), and increasing urbanisation depletes biodi-
versity in protected areas (Mcdonald et al. 2008). They
also discovered that arable land contributes to biodiver-
sity loss. Kong et al. (2021) used the human land-use
disturbance index (LDI) to assess habitat loss and frag-
mentation in global biodiversity hotspots from 1992 to
2015. Nonetheless, there is no evidence to support the
claim that human development has a negative impact on
habitats. According to the findings, developed countries
have not done a better job of conserving biodiversity
hotspots than developing countries. Despite this, rapid
urbanisation in Central Asia and the Himalaya will pose
a significant challenge to biodiversity conservation in
the future.
Similarly, Cordier et al. (2021) used hierarchical meta-
analyses to assess the effects of land-use changes such as
agriculture, cattle ranching, urbanisation, deforestation,
silviculture and selective logging on amphibian and reptile
community richness. The only land-use change that had
a significant negative impact on species richness in both
groups was urbanisation. Deforestation, silviculture and
urbanisation all had negative impacts on amphibian species
richness, while cattle raising and urbanisation had a signifi-
cant negative impact on reptile richness. However, selective
logging had no effect on the richness of either group. Hu
et al. (2021) concluded that agricultural expansion was the
primary cause of forest loss, with major losses occurring in
Sundaland, Indo-Burma and Mesoamerica. In addition, the
forest losses occurred within protected areas. As a result,
they advocated for more stringent and effective land-based
policies to prevent additional landscape fragmentation and
preserve existing species richness in the world’s biodiversity
hotspots.
In summary, the ongoing concern about rapid biodiversity
loss has fuelled an increase in academic interest in determin-
ing the possible factors that explain the extinction of various
species. We revisit the drivers of biodiversity loss in South
and Southeast Asian countries in this paper, focusing on the
land-use factors to capture habitat change. We also include
some control variables that have a significant impact on spe-
cies extinction. Furthermore, biodiversity loss measures are
extended to nine broad taxonomic groups, which include
mammals, birds, reptiles, amphibians, fishes, plants, mol-
luscs, other invertebrates and the total number of threatened
species. We use count-data estimation to generate evidence
for the causes of the region’s rapid biodiversity loss.
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63266
Data and methodology
Data
A panel dataset of 16 South and Southeast Asian coun-
tries including Bangladesh, Bhutan, Brunei, Cambodia,
India, Indonesia, Laos, Malaysia, Myanmar, Nepal, the
Philippines, Singapore, Sri Lanka, Thailand, Timor-Leste
and Vietnam from 2013 to 2018 was used. The selection
of the time periods and countries was based on the data
availability.
The threatened species can be decomposed into eight
taxonomic groups. These include the number of mammals
threatened species (MAMMAL), birds threatened species
(BIRD), reptiles threatened species (REPTILE), amphibians
threatened species (AMPHIBIAN), fish threatened species
(FISH), molluscs threatened species (MOLLUSC), other
invertebrates threatened species (OTHER) and plants threat-
ened species (PLANT). The total number of threatened spe-
cies (TOTAL) is the sum of MAMMAL, BIRD, REPTILE,
AMPHIBIAN, FISH, MOLLUSC, OTHER and PLANT.
Various versions of the IUCN Red List of Threatened
Species contain data for eight taxonomic groups of threat-
ened species. Although the IUCN Red List does not cover
all species information and is far from a worldwide species-
by-species assessment, it provides the best available data on
threatened species information and is a valuable source of
information on individual countries’ biodiversity loss status
(Clausen and York 2008; Naidoo and Adamowicz 2001).
The World Bank’s World Development Indicators
(WDI 2021) database was used to obtain the habitat change
indicator variables such as agricultural land ­(km2) (AGL),
arable land (hectares) (ARL) and forest area (­ km2) (FA), as
well as other explanatory variables such as gross domestic
product per capita (constant 2010 US$) (GDPPC) and total
population (PP). Control of corruption (CC) was obtained
from the World Bank’s Worldwide Governance Indicators
(WGI 2022) database.
The model of biodiversity loss
Land-related drivers such as land conversion, expan-
sion of land use, land-use changes, land-cover changes,
protected areas, forest cover and forest loss are largely
blamed for biodiversity threats (Bailey et al. 2016; Butch-
art et al. 2010; Cordier et al. 2021; Hu et al. 2021). Follow-
ing previous studies (Asafu-Adjaye 2003; Gren et al. 2015;
Habibullah et al. 2019; 2021), the biodiversity loss model
was specified as follows:
BL = (GDPPC, PP, HC, CC) (1)
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Environmental Science and Pollution Research (2022) 29:63260–63276
BL is measured by eight taxonomic groups of threat-
ened species: MAMMAL, BIRD, REPTILE, AMPHIB-
IAN, FISH, MOLLUSC, OTHER, PLANT and TOTAL.
GDPPC, PP, HC and CC are the regressors. GDPPC stands
for gross domestic product per capita (constant 2010 US
dollars), PP stands for total population (people) and CC
stands for control of corruption. The variable of interest
is the indicator of habitat change (HC), which is proxied
by agricultural land (AGL) (­ km2), arable land (ARL) (hec-
tares) and forest area (FA) (­ km2). The econometric model
of biodiversity loss can be specified as follows:
BLit = β0 + β1 LGDPPCit + β2 LPPit + β3 LHCit + β4 LCCit + εit
(2)
where i = 1,…, N denotes countries, t = 1,.., T denotes
time period and 𝜀 is the error term. All of the variables
were previously defined. Furthermore, all the explanatory
variables are in natural logarithm (L) to reduce the effect of
outliers on model estimates (Clausen and York 2008; Nai-
doo and Adamowicz 2001). The coefficients β1 and β2 are
expected to have positive signs. On the other hand, control of
corruption (CC) ranges from − 2.5 (the most corrupt) to + 2.5
(the least corrupt). This therefore suggests that countries
that enjoy higher level of control of corruption (low levels
of corruption) are associated with low biodiversity threats.
The expected impact of control of corruption (CC) on biodi-
versity loss is negative. In terms of habitat change influence,
agricultural land (AGL) and arable land (ARL) are expected
to have a positive impact on biodiversity loss, whereas forest
area (FA) is expected to have a negative impact on biodi-
versity loss.
Negative binomial regression models
Since the regressand (number of threatened species per year)
is count rather than continuous (Clausen and York 2008;
Naidoo and Adamowicz 2001), using the ordinary least
square (OLS) estimator on a count response variable will
result in inefficient, inconsistent and biased estimates (Hoff-
mann 2004a; Long 1997). For modelling count data, Pois-
son regression model (PRM) and negative binomial regres-
sion model (NBRM) are frequently used as alternatives to
OLS estimation. This is due to the fact that Poisson-based
regression models are intended for dependent variables in
the form of event count (Osgood 2000). Nonetheless, the
standard Poisson distribution typically assumes the follow-
ing: (i) that the conditional mean and variance are equal, also
known as equidispersion (Cupal et al. 2015; Greene 2012;
Osgood 2000; Varma et al. 2020; Winkelmann 1999), and
(ii) that the occurrences of an event (regressand) are inde-
pendent (Hoffmann 2004b).
Due to the existence of overdispersion in the data,
the NBRM is far more compelling than the PRM.
Environmental Science and Pollution Research (2022) 29:63260–63276 63267

Overdispersion occurs when the regressand’s variance Results and discussion

exceeds the mean, var(y i ) > E(y i ) (Hoffmann 2004a;
Long 1997). Typically, overdispersion is mainly due to
(i) the unobserved heterogeneity resulting from model
misspecification, omission of important predictor vari-
ables and excess zero counts (Hosseinpour et al. 2014;
Mitra and Washington 2007). (ii) Contagion (com-
monly known as state dependence), a situation char-
acterized by each event occurring, is not always an
independent event (Coxe et al. 2009). Therefore, sta-
tus of one species is dependent on the status of other
species (Shandra et al. 2009). For instance, there is
an interrelationship between animals (fishes) in the
ecosystem, indicating that big mammals (fishes) rely
on smaller mammals (fishes) for food (Clausen and
York 2008).
As a result, the negative binomial distribution for yi is
( )( )𝛼−1 ( )yi
( ) r 𝛼 −1 + yi 𝛼 −1 𝜇i
P Yi = yi ||𝜇i , 𝛼 = ( )
r 𝛼 −1 yi ! 𝛼 −1 + 𝜇i 𝛼 −1 + 𝜇i
(3)
where Y is a random variable, y i is the number of
occurrences of threatened species, r(.) is the gamma inte-
gral that specializes to a factorial for an integer argument,
α is a random parameter, µ i = exp(x iβ), x i is the vector
of independent variables, β is the vector of regression
coefficients and xiβ = β0 + β1GDPi + β2PPi + β3HCi + β4C
Ci. If the equality assumption is violated, the NBRM is
preferred over the PRM. When σ2→ 0, the NB probabil-
ity function converges to the Poisson probability func-
tion, and the log likelihood ratio test is used to assess the
validity of the equidispersion restriction (Winkelmann
and Zimmermann 1998).
The summary statistics for the key variables used in the cur-
rent study are presented in Table 1. The average total number
of threatened species in South and Southeast Asian countries
was about 502.43, with a minimum of 20 (Timor-Leste) and
a maximum of 1662 (Malaysia). Plant species suffered the
most, with the highest number of species threatened with
extinction (182.55) reported among all taxonomic groups.
Other invertebrates, fishes, mammals and birds are next,
with a mean number of threatened species of 106.07, 66.76,
50.54 and 45.81, respectively.
Molluscs, amphibians and reptiles had the fewest threat-
ened species on average, with 7.68, 19.25 and 23.60 species,
respectively. Furthermore, the variance of each threatened
species is clearly larger than its mean. This indicates that a
negative binomial regression model would be a better choice
than a Poisson regression model due to overdispersion. The
annual average GDP per capita in the 16 South and South-
east Asian countries was around $8124.05, ranging from
$664.79 (Nepal) to $59,374.40 (Singapore). On the other
hand, the average total population was 137 million people,
with a low population of 404,414 people (Brunei) and a
high population of 1400 million people (India). The agri-
cultural land (an average of 207,310.9 ­km2) and arable land
(an average of 15,185,026.0 hectares) were used in the study
to proxy habitat change indicator variables. Similarly, the
average forest area is 185,030.9 ­km2.
Table 2 shows the correlation coefficients of the regres-
sors. The majority of the explanatory variables are weakly
correlated, implying that there is little or no multicollin-
earity between them. Nonetheless, the variables of interest
(LAGL, LARL and LFA) have a strong positive correlation,

Table 1  Summary statistics N Mean Std. Dev Minimum Maximum

MAMMAL 128 50.539 41.427 4.000 192.000

BIRD 128 45.805 34.648 6.000 166.000
REPTILE 128 23.602 16.770 2.000 71.000
AMPHIBIAN 128 19.250 23.630 0.000 75.000
FISH 128 66.758 60.863 3.000 240.000
MOLLUSC 128 7.680 11.420 0.000 42.000
OTHER 128 106.070 97.228 1.000 322.000
PLANT 128 182.547 211.189 1.000 993.000
TOTAL 128 502.430 428.441 20.000 1662.000
GDPPC 128 8124.050 14,538.380 664.794 59,374.400
PP 128 137,000,000.0000 319,000,000.0000 404,414.0000 1,400,000,000.0000
AGL 96 207,310.900 436,819.100 6.600 1,800,000.000
ARL 96 15,185,026.000 38,196,422.000 560.0000 160,000,000.000
FA 96 185,030.900 262,674.600 159.3000 968,804.000
CC 112 -0.193 0.860 -1.326 2.175

13
63268
Table 2  Correlation matrix LGDPPC
LGDPPC 1.000
LPP -0.309
LAGL -0.635
LARL -0.662
LFA -0.523
LCC 0.794
with rLARL,LAGL = 0.983, rLFA,LAGL = 0.934 and rLFA,LARL =
0.898, respectively. Due to the strong correlation results, we
decided to separate the habitat change indicator variables
into separate models to avoid the problem of multicollinear-
ity (Katircioglu et al. 2019).
Table 3 presents the results of the NBRMs, whereas
Table 4 presents the average marginal effects of the
NBRMs. These tables use the NB estimator to examine the
main effects of habitat change measures extracted from the
World Development Indicator (WDI 2021): agricultural land
(LAGL), arable land (LARL) and forest area (LFA) on indi-
vidual taxonomic groups of threatened species (model 1 to
model 8) and an aggregate number of threatened species
(model 9). Meanwhile, the NBRMs include several regres-
sors, including GDP per capita (LGDPPC), population
(LPP) and corruption control (LCC). Tables 3 (panel (a))
and 4 (panel (a)) present the results for threatened species
and agricultural land, respectively, while panel (b) in Table 3
and panel (b) in Table 4 show the effects of arable land on
threatened species, and panel (c) in Table 3 and panel (c) in
Table 4 present the results for threatened species and forest
area.
The likelihood ratio test was used to evaluate the mod-
el’s goodness-of-fit. Rejection of the null hypothesis that
all parameter estimates are equal to zero implies that all
estimated models are statistically significant at the 1% level.
As a result, the nine models fit the data well in all Table 3.
Table 3 (panel (a)) has a 𝜒 2 statistic with four degrees of
freedom ranging from 57.05 to 201.70; panel (b) has a 𝜒 2
statistic ranging from 38.20 to 185.20; and panel (c) has a
𝜒 2 statistic ranging from 61.38 to 200.60.
The likelihood ratio test results show that the null hypoth-
esis that 𝛼 = 0 is strongly rejected in all models, indicating
that overdispersion exists. In all models from Table 3, the
p-values of the 𝜒 2 statistic with one degree of freedom are
less than the level of significance. This suggests that the
NBRM is better suited than the PRM. As a result, the NB
estimates are used in the following discussions. NBRMs
have previously been used in studies on biodiversity loss
(Clausen and York 2008; Mikkelson et al. 2007; Naidoo and
Adamowicz 2001).
This study distinguished between agricultural land
(LAGL), arable land (LARL) and forest area (LFA)
13
Environmental Science and Pollution Research (2022) 29:63260–63276
LPP LAGL LARL LFA LCC
-0.309 -0.635 -0.662 -0.523 0.794
1.000 0.773 0.820 0.637 -0.484
0.773 1.000 0.983 0.934 -0.718
0.820 0.983 1.000 0.898 -0.766
0.637 0.934 0.898 1.000 -0.630
-0.484 -0.718 -0.766 -0.630 1.000
measures for the effects of habitat change. Except for model
7, the coefficients of agricultural land (LAGL) and arable
land (LARL) in Table 3 (panels (a) and (b)) are positive and
significant. The findings suggest that habitat change caused
by land-use pressures, regardless of the indicators used, has
a similar direct effect on the number of threatened species
in South and Southeast Asian countries.
The average marginal effects shown in Table 4 are then
used to calculate the instantaneous rate of change in a
regressand caused by a 1-unit change in regressor (Zame-
nian et al. 2017). Both agricultural land and arable land
coefficients have statistically different effects between taxo-
nomic groups. Other things being equal, a 1% increase in
agricultural land will result in a rise of 0.022 to 1.110 total
threatened species based on the average marginal effects
of panel (a) of Table 4. Meanwhile, the average marginal
effects in Table 4, panel (b) indicate that a 1% increase in
arable land is associated with an increase in total threat-
ened species ranging from 0.024 to 1.340. Furthermore, our
findings show that the impacts of arable land expansion on
the number of threatened species in most taxonomic groups
are slightly greater than the impacts of agricultural land
development.
Threatened plants are among the eight taxonomic groups
that are most vulnerable to agricultural land and arable land
expansion. Threatened reptiles, on the other hand, are less
vulnerable to agricultural and arable land expansions. For a
more detailed discussion, average marginal effects suggest
that with a 1% increase in agricultural land and arable land,
there are approximately 0.885 and 1.152 species increases in
threatened plants, respectively. A 1% increase in agricultural
land (arable land) will result in an increase of about 0.022
(0.024) species of threatened reptiles.
According to Newbold et al. (2015), land-use changes
and related pressures reduced species richness, rarefied rich-
ness and total abundance by an average of 76.5%, 40.3%
and 39.5%, respectively, in the worst-affected habitats.
These changes resulted in a reduction of 13.6% (species
richness), 10.7% (total abundance) and 8.1% (rarefied rich-
ness) in local terrestrial biodiversity by 2005. Similarly,
Perrings and Halkos (2015) found that a 1% increase in
agricultural land conversion leads to a 2.2–3.7% increase in
threatened endemic mammals, birds and plants. Naidoo and
Environmental Science and Pollution Research (2022) 29:63260–63276 63269

Table 3  Negative binomial regression results

(a) Agricultural land on threatened species

(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIB- FISH MOLLUSC OTHER PLANT TOTAL
IAN
LGDPPC 0.447*** 0.306*** 0.374*** 1.222*** 0.713*** 1.110*** 1.426*** 1.342*** 0.854***
(8.35) (5.82) (5.78) (5.85) (9.17) (3.91) (7.86) (13.73) (14.98)
LPP − 0.00613 0.137*** 0.223*** − 0.0158 0.325*** − 0.723*** 0.639*** 0.111** 0.208***
(− 0.20) (4.55) (6.27) (− 0.12) (8.22) (− 2.58) (6.91) (2.18) (6.98)
***
LAGL 0.280 0.173*** 0.0988*** 0.638*** 0.0878** 1.306*** − 0.0373 0.432*** 0.220***
(10.00) (6.48) (3.11) (4.84) (2.52) (3.96) (− 0.56) (9.23) (8.46)
LCC − 0.215 0.0653 − 1.331*** − 1.838** − 2.052*** − 3.477*** − 3.474*** − 0.554 − 0.708***
(− 0.96) (0.28) (− 4.35) (− 2.22) (− 5.96) (− 2.70) (− 4.69) (− 1.49) (− 3.19)
C − 2.657*** − 2.949*** − 5.077*** − 13.97*** − 8.625*** − 9.509*** − 18.30*** − 12.62*** − 6.915***
(− 5.15) (− 5.84) (− 7.37) (− 6.77) (− 11.31) (− 2.95) (− 11.15) (− 13.08) (− 13.07)
Lɑ − 2.330 − 2.442 − 2.681 0.00971 − 1.873 0.456 − 0.369 − 1.163 − 2.155
LR(1) 404.12 259.19 57.41 1040.13 316.86 467.01 2465.54 3558.30 3754.19
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
LR(4) 143.9 146.5 162.2 82.63 170.4 57.05 111.3 163.0 201.7 (0.000)
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
2
R 0.156 0.164 0.208 0.113 0.173 0.108 0.105 0.139 0.147
N 96 96 96 96 96 96 96 96 96
LL − 390.3 − 374.3 − 308.1 − 323.9 − 407.3 − 234.9 − 474.6 − 504.2 − 586.4
(b) Arable land on threatened species
(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIB- FISH MOLLUSC OTHER PLANT TOTAL
IAN
LGDPPC 0.500*** 0.325*** 0.393*** 1.327*** 0.748*** 1.491*** 1.390*** 1.452*** 0.892***
(8.54) (5.76) (5.87) (5.74) (9.46) (4.67) (7.36) (12.63) (13.56)
LPP − 0.0836** 0.102*** 0.212*** − 0.0746 0.268*** − 0.230 0.683*** 0.0108 0.162***
(− 2.08) (2.66) (4.67) (− 0.42) (5.34) (− 0.90) (5.88) (0.14) (3.88)
LARL 0.353*** 0.202*** 0.106** 0.684*** 0.141*** 0.715*** − 0.0815 0.534*** 0.262***
(9.21) (5.63) (2.53) (3.84) (3.07) (2.73) (− 0.84) (7.16) (6.65)
LCC 0.154 0.255 − 1.272*** − 1.269 − 1.823*** − 3.106* − 3.634*** − 0.0825 − 0.480*
(0.62) (1.02) (− 3.82) (− 1.41) (− 5.04) (− 1.89) (− 4.62) (− 0.19) (− 1.90)
C − 3.850*** − 3.582*** − 5.519*** − 16.83*** − 9.014*** − 17.10*** − 18.01*** − 14.86*** − 7.851***
(− 6.86) (− 6.63) (− 8.01) (− 7.52) (− 12.02) (− 5.04) (− 10.81) (− 13.18) (− 12.94)
Lɑ − 2.204 − 2.330 − 2.607 0.119 − 1.913 0.688 − 0.375 − 0.951 − 1.973
LR(1) 525.66 311.97 63.53 1156.15 302.04 647.60 2393.64 6108.02 5808.50
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
LR(4) 136.7 139.1 159.0 71.17 173.1 38.20 111.7 144.7 185.2
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
R2 0.148 0.155 0.204 0.0974 0.176 0.0725 0.105 0.124 0.135
N 96 96 96 96 96 96 96 96 96
LL − 393.8 − 378.0 − 309.7 − 329.7 − 406.0 − 244.4 − 474.4 − 513.3 − 594.7
(c) Forest area on threatened species
(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIB- FISH MOLLUSC OTHER PLANT TOTAL
IAN
LGDPPC 0.309*** 0.230*** 0.333*** 0.914*** 0.680*** 0.702*** 1.427*** 1.146*** 0.760***
(8.20) (4.99) (5.26) (4.41) (9.64) (2.80) (8.03) (11.49) (14.26)
LPP 0.0850*** 0.194*** 0.252*** 0.436*** 0.328*** − 0.0951 0.636*** 0.319*** 0.294***
(4.89) (9.16) (10.23) (4.41) (12.23) (− 0.83) (7.98) (6.58) (12.13)

13
63270 Environmental Science and Pollution Research (2022) 29:63260–63276

Table 3  (continued)

LFA 0.313*** 0.196*** 0.123*** 0.224* 0.153*** 1.348*** − 0.0542 0.345*** 0.222***
(15.73) (8.27) (4.33) (1.84) (4.81) (6.77) (− 0.74) (6.12) (8.22)
LCC − 0.0646 0.124 − 1.260*** − 2.140** − 1.800*** − 1.639 − 3.475*** − 0.674* − 0.699***
(− 0.38) (0.58) (− 4.30) (− 2.39) (− 5.68) (− 1.39) (− 4.77) (− 1.66) (− 3.13)
C − 3.656*** − 3.683*** − 5.559*** − 14.97*** − 9.165*** − 18.19*** − 18.06*** − 13.89*** − 7.790***
(− 9.26) (− 7.65) (− 8.36) (− 6.60) (− 12.87) (− 6.38) (− 10.89) (− 12.31) (− 13.97)
Lɑ − 3.209 − 2.678 − 2.782 0.178 − 2.086 0.153 − 0.372 − 0.902 − 2.133
LR(1) 130.02 190.37 54.55 1142.94 231.68 441.76 2465.41 4490.96 3994.52
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
LR(4) 200.6 164.3 171.0 61.38 183.9 86.81 111.6 138.8 200.2
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
R2 0.217 0.184 0.220 0.0840 0.187 0.165 0.105 0.118 0.146
N 96 96 96 96 96 96 96 96 96
LL − 361.9 − 365.4 − 303.7 − 334.6 − 400.5 − 220.1 − 474.4 − 516.3 − 587.2

t statistics in parentheses. *, ** and *** denote p < 0.1, p < 0.05 and p < 0.01, respectively. L is the natural logarithm, Lɑ is the log-transformed
alpha, LR(1) is the likelihood ratio test of ɑ = 0 with one degree of freedom, LR(4) is the likelihood ratio test that all the regression coefficients
are zero with four degrees of freedom, R2 is the pseudo R2 and LL is the log likelihood

Adamowicz’s (2001) findings also indicated that land-use
variables appear to have a significant effect on the number
of threatened species in 5 of 21 NBRMs. In particular, an
increase in protected areas reduces the number of threatened
amphibians (in two models), whereas more domesticated
land (for agricultural use) has higher levels of threatened
invertebrates (in two models), and there is a nonlinear rela-
tionship between forest area and the number of threatened
mammal species (in one model). According to a recent study
by Tommasi et al. (2021), agricultural cover expansion is
likely to reduce plant and pollinator richness by simplifying
the landscape with less habitat, whereas land-use changes
have a significant impact on amphibian and reptile richness
(Cordier et al. 2021).
Nonetheless, the parameter estimates for forest area
(LFA) in Table 3 (panel (c)) are significant in most mod-
els, but of an unexpected sign. Despite significant efforts,
biodiversity loss remains significant, according to Butch-
art et al. (2010). These efforts include local successes,
for example, continuous monitoring of protected areas,
the sustainability of forest management, invasive spe-
cies policies and biodiversity-related assistance. Gren
et al. (2015), on the other hand, presented some intrigu-
ing findings regarding land-use variables. Arable land
had an unexpectedly negative and significant effect on all
threatened species and five taxonomic groups of threat-
ened species (mammals, birds, amphibians, plants and
reptiles). Another finding from Gren et al. (2015) was
that protected areas have yielded inconclusive results.
More protected area has a negligible, albeit positive,
impact on all threatened species; however, protected
area has a negative impact on mammals, birds and rep-
tiles and a positive impact on amphibians and plants. One
possible explanation for arable land’s negative impact is
that nations with a relatively large arable land area are
associated with a large number of species. Another, possi-
bly valid, reason is the extensive or intensive management
of arable land use.
In contrast to other studies, some studies found no statisti-
cal evidence that agricultural land (Kerr and Currie 1995),
protected areas (Kerr and Burkey, 2002) or arable land (Li
et al. 2013) posed more threats to biodiversity. Kerr and
Currie (1995), for example, discovered that habitat loss (as
measured by agricultural land) is not a significant threat to
bird species. Kerr and Burkey (2002) found that protected
areas have no effect on the number of threatened or endemic
bird and mammal species. At the global level, no correlation
was found between threatened mammals and protection, but
at the subecoregional level in California, a positive correla-
tion was found between the number of threatened plants and
protection (Underwood et al. 2009).
Other anthropogenic and socioeconomic activities, such
as per capita income and population size, have a signifi-
cant impact on species at risk of extinction. Kerr and Cur-
rie (1995) discovered that the former was closely related to
threatened mammals, whereas the latter appears to affect
the number of threatened bird species. In panel fixed effect
regression, Bhuiyan et al. (2018) found that GDP per capita
and forest area are positively related, whereas population
growth has no effect on forest area. GDP per capita has con-
sistently positive effects and is statistically significant ( p <
0.01) in all models for all tables in this study. A high level
of per capita income is associated with a greater number
of threatened species. Based on average marginal effects,
a 1% increase in per capita income raises the number of
threatened species in South and Southeast Asian countries

13
Environmental Science and Pollution Research (2022) 29:63260–63276 63271

Table 4  Marginal effects of negative binomial regression results

(a) Agricultural land on threatened species

(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIBIAN FISH MOLLUSC OTHER PLANT TOTAL
LGDPPC 22.53*** 13.53*** 8.511*** 33.23*** 43.97*** 10.86** 169.0*** 274.7*** 431.4***
(7.39) (5.53) (5.49) (3.13) (7.67) (2.37) (5.12) (6.69) (10.57)
LPP − 0.309 6.058*** 5.083*** − 0.431 20.00*** − 7.078* 75.69*** 22.68** 105.2***
(− 0.20) (4.41) (5.94) (− 0.12) (7.20) (− 1.96) (4.63) (2.17) (6.48)
LAGL 14.10*** 7.658*** 2.247*** 17.36*** 5.409** 12.78** − 4.416 88.45*** 111.0***
(8.53) (6.03) (3.05) (2.87) (2.46) (2.36) (− 0.56) (5.59) (7.06)
LCC − 10.82 2.888 − 30.27*** − 49.99** − 126.5*** − 34.03** − 411.8*** − 113.4 − 357.6***
(− 0.96) (0.28) (− 4.27) (− 2.00) (− 5.58) (− 1.98) (− 4.11) (− 1.47) (− 3.16)
N 96 96 96 96 96 96 96 96 96
(b) Arable land on threatened species
(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIBIAN FISH MOLLUSC OTHER PLANT TOTAL
LGDPPC 25.59*** 14.46*** 8.977*** 39.62*** 46.39*** 16.48** 162.2*** 313.6*** 457.0***
(7.40) (5.45) (5.54) (2.82) (7.78) (2.21) (4.90) (5.94) (9.52)
LPP − 4.277** 4.559*** 4.840*** − 2.226 16.63*** − 2.542 79.71*** 2.331 82.94***
(− 2.05) (2.64) (4.56) (− 0.41) (5.10) (− 0.81) (4.48) (0.14) (3.84)
*** ***
LARL 18.05 9.012 2.419** 20.41** 8.752*** 7.909* − 9.510 115.2*** 134.0***
(7.75) (5.25) (2.48) (2.25) (2.94) (1.69) (− 0.85) (4.50) (5.65)
LCC 7.902 11.34 − 29.04*** − 37.89 − 113.2*** − 34.35 − 424.2*** − 17.82 − 246.0*
(0.62) (1.01) (− 3.78) (− 1.39) (− 4.84) (− 1.62) (− 4.18) (− 0.19) (− 1.91)
N 96 96 96 96 96 96 96 96 96
(c) Forest area on threatened species
(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIBIAN FISH MOLLUSC OTHER PLANT TOTAL
LGDPPC 15.47*** 10.22*** 7.603*** 21.07*** 42.31*** 9.231** 168.4*** 228.2*** 386.1***
(7.77) (4.85) (5.06) (3.02) (8.12) (2.15) (5.25) (6.12) (10.45)
LPP 4.253*** 8.624*** 5.764*** 10.05*** 20.42*** − 1.250 75.10*** 63.55*** 149.5***
(4.77) (8.21) (8.98) (3.22) (9.59) (− 0.84) (4.82) (4.98) (9.44)
LFA 15.67*** 8.685*** 2.805*** 5.156 9.526*** 17.73** − 6.402 68.78*** 112.7***
(13.30) (7.51) (4.18) (1.54) (4.47) (2.56) (− 0.75) (4.30) (6.83)
LCC − 3.233 5.486 − 28.80*** − 49.35** − 112.1*** − 21.55 − 410.2*** − 134.4 − 355.0***
(− 0.38) (0.58) (− 4.23) (− 2.14) (− 5.41) (− 1.31) (− 4.16) (− 1.64) (− 3.12)
N 96 96 96 96 96 96 96 96 96

t statistics in parentheses. *, ** and *** denote p < 0.1, p < 0.05 and p < 0.01, respectively

by about 0.085 to 4.314 species in Table 4 (panel (a)); 0.090
to 4.570 species in panel (b); and 0.076 to 3.861 species in
panel (c).
In 19 of the 27 models (Table 3 from panel (a) to
panel (c)), the coefficient for total population (LPP) has
a positive and significant effect. This suggests that rapid
population growth is more likely to exacerbate species
threats and threaten biodiversity loss. Similar positive
impact of population growth on threatened species was
found by Habibullah et al. (2016). According to Asafu-
Adjaye (2003), increased population causes habitat loss
due to increased demand for human needs such as food,
shelter and other services. The expanding human popu-
lation may endanger tropical forests through alternative
human activities such as agricultural expansion, hunting,
logging and other human settlement (Giam 2017). Crop-
per and Griffiths (1994) show that in Africa, every 100
persons per 1000 hectares increase in rural population
density reduces deforestation by about 0.33 percentage
points, whereas each 100 persons increase in population
density raises the threat to mammal species by about one
percentage point and the threat to bird species by about 0.5
percentage points in model 2 and model 3 (McPherson and
Nieswiadomy 2005).

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63272 Environmental Science and Pollution Research (2022) 29:63260–63276

Table 5  Poisson regression results

(a) Agricultural land on threatened specie

(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIBIAN FISH MOLLUSC OTHER PLANT TOTAL
LGDPPC 0.405*** 0.317*** 0.269*** 0.431*** 0.540*** 0.832*** 1.116*** 0.884*** 0.684***
(17.34) (12.17) (6.74) (10.44) (20.47) (13.48) (45.77) (65.23) (81.30)
LPP 0.0156 0.145*** 0.210*** 0.202*** 0.300*** − 0.581*** 0.420*** 0.0600*** 0.169***
(0.98) (9.19) (8.49) (7.75) (18.57) (− 8.13) (34.84) (7.69) (35.80)
LAGL 0.268*** 0.158*** 0.0900*** 0.266*** 0.139*** 0.832*** 0.00484 0.374*** 0.210***
(18.21) (11.24) (4.03) (11.32) (10.09) (11.54) (0.53) (57.03) (53.48)
LCC − 0.222** − 0.176 − 0.994*** 0.541*** − 0.940*** − 1.057*** − 2.241*** 0.646*** − 0.187***
(− 1.99) (− 1.40) (− 4.88) (2.73) (− 6.87) (− 3.72) (− 17.65) (10.20) (− 4.53)
C − 2.576*** − 3.042*** − 3.863*** − 7.005*** − 7.214*** − 4.058*** − 12.21*** − 7.239*** − 4.683***
(− 9.96) (− 10.54) (− 8.45) (− 14.10) (− 21.63) (− 5.65) (− 38.61) (− 43.34) (− 46.15)
LR(4) 1866.7 (0.000) 1549.8 (0.000) 831.0 1415.3 4016.6 (0.000) 516.0 6852.3 (0.000) 14,226.0 (0.000) 27,026.7
(prob χ2) (0.000) (0.000) (0.000) (0.000)
R2 0.612 0.606 0.552 0.456 0.780 0.355 0.667 0.757 0.846
N 96 96 96 96 96 96 96 96 96
LL − 592.3 − 503.9 − 336.8 − 844.0 − 565.7 − 468.4 − 1707.3 − 2283.3 − 2463.5
(b) Arable land on threatened species
(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIBIAN FISH MOLLUSC OTHER PLANT TOTAL
LGDPPC 0.445*** 0.341*** 0.286*** 0.488*** 0.569*** 0.850*** 1.201*** 0.945*** 0.733***
(18.71) (12.80) (7.11) (10.73) (22.31) (12.75) (47.22) (63.40) (82.17)
LPP 0.00411 0.146*** 0.222*** 0.336*** 0.241*** − 0.118** 0.550*** 0.138*** 0.218***
(0.23) (7.61) (7.39) (10.19) (12.30) (− 2.30) (33.00) (14.12) (35.54)
LARL 0.262*** 0.145*** 0.0721*** 0.118*** 0.183*** 0.342*** − 0.111*** 0.270*** 0.150***
(16.22) (8.62) (2.74) (4.33) (11.13) (7.49) (− 8.41) (31.66) (28.80)
LCC − 0.0306 − 0.166 − 1.039*** − 0.105 − 0.682*** − 0.966*** − 3.330*** 0.121 − 0.561***
(− 0.25) (− 1.21) (− 4.68) (− 0.42) (− 4.71) (− 2.58) (− 21.63) (1.50) (− 11.23)
C − 3.617*** − 3.684*** − 4.304*** − 8.712*** − 7.585*** − 8.068*** − 13.65*** − 9.034*** − 5.909***
(− 14.18) (− 12.83) (− 9.68) (− 15.88) (− 24.38) (− 10.62) (− 41.66) (− 49.45) (− 55.22)
LR(4) 1738.0 (0.000) 1489.7 (0.000) 821.7 1287.8 4034.0 (0.000) 316.6 6924.6 11,658.0 (0.000) 24,955.8
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000)
R2 0.570 0.582 0.546 0.415 0.784 0.218 0.674 0.620 0.781
N 96 96 96 96 96 96 96 96 96
LL − 656.7 − 534.0 − 341.4 − 907.8 − 557.0 − 568.2 − 1671.2 − 3567.3 − 3498.9
(c) Forest area on threatened species
(1) (2) (3) (4) (5) (6) (7) (8) (9)
Variables MAMMAL BIRD REPTILE AMPHIBIAN FISH MOLLUSC OTHER PLANT TOTAL
LGDPPC 0.310*** 0.259*** 0.238*** 0.437*** 0.484*** 0.666*** 1.130*** 0.787*** 0.633***
(13.17) (9.86) (5.84) (9.16) (18.01) (10.30) (45.13) (54.04) (71.42)
LPP 0.0742*** 0.186*** 0.240*** 0.403*** 0.330*** − 0.204*** 0.430*** 0.210*** 0.250***
(6.73) (16.37) (14.58) (20.63) (29.55) (− 5.58) (45.16) (33.71) (67.50)
LFA 0.340*** 0.187*** 0.0972*** 0.0906*** 0.166*** 0.730*** − 0.00688 0.327*** 0.197***
(24.77) (14.34) (5.24) (4.64) (13.91) (14.53) (− 0.80) (49.53) (50.63)
LCC − 0.105 − 0.153 − 1.007*** − 0.309 − 0.916*** − 0.781*** − 2.351*** 0.183*** − 0.430***
(− 0.96) (− 1.24) (− 5.12) (− 1.39) (− 7.10) (− 2.59) (− 20.64) (2.73) (− 10.30)
C − 3.783*** − 3.700*** − 4.259*** − 8.795*** − 7.662*** − 8.512*** − 12.39*** − 8.765*** − 5.658***
(− 15.37) (− 13.33) (− 9.96) (− 16.12) (− 25.35) (− 12.56) (− 41.94) (− 51.12) (− 56.30)
LR(4) 2197.5 1636.5 842.6 1291.2 4115.3 571.0 6852.6 (0.000) 13,269.1 26,784.8
(prob χ2) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000) (0.000)
R2 0.720 0.640 0.560 0.416 0.799 0.393 0.667 0.706 0.838
N 96 96 96 96 96 96 96 96 96
LL − 426.9 − 460.6 − 331.0 − 906.0 − 516.4 − 440.9 − 1707.1 − 2761.7 − 2584.4

t statistics in parentheses. *, ** and *** denote p < 0.1, p < 0.05 and p < 0.01, respectively. LR(4) is the likelihood ratio test that all the regression
coefficients are zero with four degrees of freedom, R2 is the pseudo R2 and LL is the log likelihood

13
Environmental Science and Pollution Research (2022) 29:63260–63276
Finally, in Tables 3, 4 and 5, the control of corruption
(LCC) appears to have a negative and significant (p < 0.01)
effect on all types of threatened species. This suggests that
good governance can help to reduce biodiversity loss if cor-
ruption is kept under control. These findings are consistent
with those of other studies (Dietz and Adger 2003; Gren
et al. 2015; Pandit and Laband 2009). Pandit and Laband
(2009) discovered a link between the extinction of mam-
mals and vascular plant species and economic freedom.
They also discovered evidence of the effects of corruption
on the extinction of birds and reptiles. Similarly, Dietz and
Adger (2003) found that democracy has a significant impact
on species richness in both linear and hyperbolic equations.
Gren et al. (2015) discovered that a certain level of institu-
tional capacity (as measured by corruption and democracy)
is required before further increases in economic growth will
result in a decrease in threatened species.
Robustness checks
We report the Poisson regression results using the same
explanatory variables in the robustness checks. In agree-
ment with Table 3, all models in Table 5 produce similar
results. Table 5, panels (a) and (b), show that agricultural
land (LAGL) and arable land (LARL) remain positive and
significant with p < 0.01 in all models except model 7,
indicating that the land-use effect exists in the majority of
threatened species. These findings are robust regardless of
the habitat change measures and estimations used.
However, with the exception of model 7, another land-use
effect—forest area (LFA)—has unexpectedly positive and
significant impacts on the number of species threatened. In
most models, the parameter estimates for income per capita
(LGDPPC) and total population (LPP) variables have a posi-
tive and significant influence on the number of threatened
species. As a result, it is noteworthy to suggest that biodi-
versity loss is responsive to anthropogenic factors. Finally,
reducing corruption has the potential to reduce threats to
species. Overall, the results of Table 5 are entirely consistent
with those of Table 3.
Conclusions
Biodiversity loss appears to be the most serious environmen-
tal concern confronting the world today. South and Southeast
Asia outnumber other Asian regions in terms of the number
of threatened species encountered between 2013 and 2018.
Furthermore, some South and Southeast Asian countries
experienced a significant decline in biodiversity richness
for a period of time. Most biodiversity hotspots are under
threat from habitat change, which may be the primary driver.
63273
Previous research has found that the effects of habitat change
are closely related to land-use changes.
This study adds to the body of knowledge about biodi-
versity decline due to habitat change by incorporating three
land-use indicator variables: agricultural land, arable land
and forest land, as well as several anthropogenic and socio-
economic factors. Furthermore, developing countries have
experienced severe biodiversity declines, particularly in
South and Southeast Asia, where much of the biodiversity
is concentrated, so we have limited this study to the coun-
tries in this region. This study also provides a more in-depth
examination of the biodiversity loss indicator, which is used
as a dependent variable. This includes mammals, birds, rep-
tiles, amphibians, fish, molluscs, other invertebrates, plants
threatened by extinction and total threatened species. As a
result, the current study seeks to contribute to the existing
literature on biodiversity loss by investigating habitat change
as one of the major drivers of biodiversity loss in South and
Southeast Asian countries from 2013 to 2018.
Poisson-based regression models are intended for mod-
elling count data with counts of events as the regressand.
However, the negative binomial regression models may be
the best estimation option because it can account for over
dispersion. Furthermore, regardless of the types of threat-
ened species or habitat change measures, the Wald test indi-
cates the goodness-of-fit for the estimated models. Moreo-
ver, the likelihood ratio tests show statistically significant
overdispersion in the estimated models.
Using different taxonomic groups to characterize the
response of individual threatened species, it was discovered
that threatened species increased with an increase in agricul-
tural and arable land. Nonetheless, the effects of arable land
on threatened species are slightly greater than those of agri-
cultural land. Among taxonomic groups, threatened plants
outnumber other types of threatened species as a result of
agricultural land and arable land expansion, whereas threat-
ened reptiles are among the less vulnerable to agricultural
land and arable land expansion. Forest area is another factor
that contributes to a higher number of threatened species.
Surprisingly, the number of threatened species
increased as available forest area increased. Aside from
habitat change, other anthropogenic and socioeconomic
factors are important drivers of biodiversity loss. In South
and Southeast Asian countries, income per capita and total
population had a significant and positive effect on the
number of threatened species. Finally, good governance,
as represented by high levels of corruption control, can
help to reduce biodiversity loss. In terms of robustness,
the empirical results appear to be robust across different
types of threatened species, habitat change measures and
Poisson-based regressions.
On top of that, this study employs the panel data, and the
inference is for regional perspectives as overall findings and
13
63274
not for a specific country conclusion. This is because South
and Southeast Asian countries consist of 16 countries of
Bangladesh, Bhutan, Brunei, Cambodia, India, Indonesia,
Laos, Malaysia, Myanmar, Nepal, the Philippines, Singa-
pore, Sri Lanka, Thailand, Timor-Leste and Vietnam. These
countries are composed of two blocks of regional coopera-
tion: the South Asian Association for Regional Coopera-
tion (SAARC)—the regional intergovernmental organization
and geopolitical union of states in South Asia that comprise
Bangladesh, Bhutan, India, Nepal and Sri Lanka, while
the ASEAN (The Association of Southeast Asian Nations)
consist of Brunei, Cambodia, Indonesia, Laos, Malaysia,
Myanmar, the Philippines, Singapore, Thailand, Timor-
Leste and Vietnam. These two blocks are in itself is not fully
integrated, more so if to collectively cooperate on a single
policy, say on forest or biodiversity conservation.
Several important policy implications can be derived
from this study. First, in pursuing higher level of economic
development and income, the government should increase
spending on education and campaign to educate and promote
the population to be more environmentally friendly. For
example, promoting biodiversity abundance in a destination
country can be used to enhance growth in the tourism sector,
by luring both domestic and international tourists. Second,
increase in the conservation of sustainable green area and/
or national forest park would be able to protect the habitat
of various species of the flora and fauna. Loss of habitat
due to deforestation and increase in agricultural activities
will reduce the reproductive rate of the flora and fauna and
force them into extinction. Lastly, government of weak and
poor governance would like to increase the loss of biodiver-
sity. Corrupt officials in the public sectors are more likely
to allow illegal logging, poaching and other illegal activities
that could further degrade the environment and biodiversity
loss. In such a case, the government should increase its effort
to eliminate or control corruption among its officials, as well
as extent good governance practices to the private sector.
Author contribution Jen-Eem Chen: introduction and review of the
literature. Thian-Hee Yiew: data collection and estimation method.
Yan-Ling Tan: abstract, introduction, results and discussion, as well as
paper conclusion. Muzafar Shah Habibullah: editing and proofreading.
The final manuscript was read and approved by all authors.
Funding The Universiti Teknologi MARA Cawangan Johor Kampus
Segamat funded this research through the Bestari Research Grant Phase
1/2021 (Grant Number: 600-UiTMCJ (PJIA.5/2)).
Data availability The information in this paper comes from the IUCN
Red List of Threatened Species (various versions), the Worldwide
Governance Indicators database and the World Development Indica-
tor (WDI) database. On reasonable request, it is available from the
corresponding author.
13
Environmental Science and Pollution Research (2022) 29:63260–63276
Declarations
Ethics approval and consent to participate Not applicable.
Consent for publication Not applicable.
Competing interests The authors declare no competing interests.
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