!

Open Veterinary Journal, (2026), Vol. 16(8): 5096–5113

Review Article

10.5455/OVJ.2026.v16.i8.6

---

Transboundary expansion of the foot-and-mouth disease virus serotype SAT1 and its implications for Indonesia’s biosecurity and disease prevention strategies

Harimurti Nuradji^1*, Aswin Rafif Khairullah¹, Ni Luh Putu Indi Dharmayanti¹, Rahmat Setya Adji¹, Agus Wiyono¹, Susan Maphilindawati Noor¹, Fitrine Ekawasti¹, Diana Nurjanah¹, Muharam Saepulloh¹, Indrawati Sendow¹, Muhammad Khaliim Jati Kusala¹, Ima Fauziah¹, Syahputra Wibowo², Ririn Setyowati³, Neny Santy Jelita Lumbantoruan⁴, Eka Oktarianti⁵, Rara Ayu Andinie⁶ and Arundhina Girishanta⁷

¹Research Center for Veterinary Science, National Research and Innovation Agency (BRISN), Bogor, Indonesia

²Eijkman Research Center for Molecular Biology, National Research and Innovation Agency (BRIN), Bogor, Indonesia

³Indonesian Quarantine Authority, Jakarta, Indonesia

⁴PT. Juang Jaya Abdi Alam, Lampung, Indonesia

⁵Animal Husbandry and Animal Health Services, West Sumatera, Indonesia

⁶Faculty of Mathematics and Natural Sciences, Program Study of Biology, Universitas Sriwijaya, Palembang, Indonesia

⁷UPTD RPH Dinas Ketahanan Pangan dan Pertanian, Indramayu, Indonesia

*Corresponding Author: Harimurti Nuradji. Research Center for Veterinary Science, Research Organization for Health, National Research and Innovation Agency (BRIN), Bogor, Indonesia. Email: hari056 [at] brin.go.id

Submitted: 30/04/2026 Revised: 07/07/2026 Accepted: 16/07/2026 Published: 08/08/2026

---

Abstract

Foot-and-mouth disease (FMD) is a highly contagious viral disease that affects cloven-hoofed animals and causes major economic losses in the livestock sector. Southern African Territories 1 (SAT1) is an important FMD serotype that was previously restricted to Sub-Saharan Africa but has recently shown evidence of wider geographic spread. Molecular surveillance and phylogenetic studies indicate that SAT serotypes may spread through international livestock trade and animal movement beyond endemic regions. Among SAT serotypes, SAT1 is of particular concern because of its high antigenic diversity, extensive genetic variability, and limited cross-protection with existing vaccines. These factors raise concerns regarding their potential introduction into disease-free regions, such as Indonesia. This review analyzes the virological and epidemiological characteristics of FMDV SAT1 and evaluates its possible introduction routes into Indonesia. The major pathways include the movement of animals and animal products, human mobility, global trade, and environmental transmission. The vulnerability of Indonesia was assessed based on livestock production systems, previous FMD outbreaks, immunity gaps, and the capacity of biosecurity and surveillance systems. The findings suggest that viral characteristics, increasing international trade, and uneven implementation of biosecurity and vaccination programs influence the risk of SAT1 introduction. Although precise quantitative estimation remains difficult due to limited surveillance data, transboundary animal movement and global trade increase the likelihood of SAT1 introduction into Southeast Asia. Strengthening farm-level biosecurity, improving genomic surveillance, optimizing vaccination strategies, and establishing vaccine banks containing antigenically matched SAT1 strains are essential for improving preparedness and emergency response capacity.

Keywords: Biosecurity, Foot-and-mouth disease, Indonesia, SAT1, Vaccination.

---

Introduction

Foot-and-mouth disease (FMD) is an infectious animal disease that has the most significant economic impact on the global livestock sector (Khairullah et al., 2024). The FMD virus (FMDV), a member of the genus Aphthovirus and the family Picornaviridae, causes this illness (Rahman et al., 2025). It is highly contagious and can cause significant losses due to reduced livestock productivity, trade restrictions, and disruption of the animal food supply chain stability (Mohamadin et al., 2025). The primary feature of FMDV is the presence of seven immunologically distinct serotypes: O, A, C, Asia1, Southern African Territories 1 (SAT1), SAT2, and SAT3 (Mabunda et al., 2026). These serotypes do not exhibit effective cross-protection, making vaccination-based control difficult and heavily reliant on the antigenic match of the vaccine strain to the field virus (Jiang et al., 2025).

The SAT1 serotype, which has historically been limited to the Sub-Saharan African region and has an endemicity pattern sustained by interactions between domestic livestock and wildlife reservoirs, particularly the African buffalo (Syncerus caffer), is particularly significant from an epidemiological standpoint (Duchatel et al., 2019). Nonetheless, signs of shifting regional distribution patterns have been observed over the past 10 years, which point to possible spread outside of the conventional endemic zones (Vosloo et al., 2002). Recent molecular characterization studies based on VP1 sequencing and phylogenetic analysis have demonstrated genetic relationships between transboundary SAT outbreaks and East African lineages, highlighting the capacity of SAT serotypes to spread through international livestock movement and trade-associated pathways (Gizaw et al., 2025). This phenomenon has caused alarm on a global scale because it shows a growing danger of transboundary transmission, which can be impacted by the dynamics of international trade, animal movement, and flaws in animal health surveillance systems in various countries (An et al., 2025).

The evolving FMDV epidemiological environment, including the potential spread of SAT1, highlights the necessity for risk assessment in previously unaffected nations (Humphreys et al., 2025). Indonesia has a sizable population of ruminant livestock and a smallholder livestock-dominated livestock system, making it fundamentally vulnerable to quickly spreading infectious illnesses (Rehman et al., 2025). Although no confirmed endemic circulation of SAT1 has yet been reported in Southeast Asia, increasing connectivity through trade, transportation, and distribution of animal products continues to increase the theoretical risk of SAT1 introduction into Indonesia and neighboring countries (Ahmed et al., 2026). Furthermore, the geographic location of Indonesia as an archipelagic nation with significant animal and animal product trade makes the risk of cross-border disease introduction more problematic (Adnyana et al., 2023). Nevertheless, the national livestock system is susceptible to the introduction of novel infectious agents and the resurgence of long-undetected diseases, as evidenced by epidemiological experience with the foot-and-mouth disease (FMD) outbreak in 2022 (Sutawi et al., 2023).

Therefore, improving national readiness requires an understanding of the virological traits of SAT1, the dynamics of its propagation, and the risk pathways of its introduction. Gaps in the surveillance system, vaccination effectiveness, and biosecurity capacity must be evaluated to find weak places in the disease control system (Elrashedy et al., 2025a). Furthermore, the integration of the One Health approach—which connects animal, human, and environmental health—becomes increasingly important to combat complex and multivariate transboundary disease risks (Danasekaran, 2024).

Therefore, this review article aims to (i) analyze the biological and epidemiological characteristics of FMDV serotype SAT1, (ii) examine global distribution patterns and their driving factors, (iii) evaluate potential routes of introduction into Indonesia, and (iv) formulate implications for evidence-based prevention and control strategies. This research is anticipated to reinforce national biosecurity regulations and improve readiness for potential FMDV threats.

Virology and epidemiology of FMDV SAT1 infection

The FMDV serotype SAT1 belongs to the genus Aphthovirus. Because of its complex epidemiological features, including high genetic variation, little cross-protection, and dynamic geographic distribution patterns, extensive virological and epidemiological research is required to understand the mechanisms underlying its persistence and spread.

Characteristics of the viruses

FMDV is a positive single-stranded RNA virus (+ssRNA) that is a member of the Picornaviridae family and genus Aphthovirus (Gao et al., 2016). The single lengthy open reading frame that makes up the FMDV genome, which is between 8.3 and 8.5 kb in size, codes for a polyprotein that is then broken down into structural and non-structural proteins by proteases (Ekanayaka et al., 2021). A non-enveloped icosahedral capsid is formed by the primary structural proteins VP1, VP2, VP3, and VP4. VP1 is essential for attaching to host cell receptors via the arginine-glycine-aspartic acid motif (Li et al., 2021).

The mutation rate of the RNA-dependent RNA polymerase, which lacks a proofreading mechanism, is the primary cause of the exceptionally high level of genetic and antigenic variation found in FMDV (Li et al., 2020). Seven different immunological serotypes—O, A, C, Asia 1, SAT1, SAT2, and SAT3—are produced as a result of this variability and do not offer significant cross-protection (Zewdie et al., 2023). Among these serotypes, SAT1 exhibits considerable genetic variability and various geographically dispersed topotypes, particularly in Sub-Saharan Africa (Sangula et al., 2010). This review focuses on SAT1 because its substantial antigenic heterogeneity and rapid evolutionary dynamics present major challenges for vaccine matching and outbreak preparedness. Nevertheless, SAT2 and SAT3 should also be recognized as potential threats to Asian countries because all SAT serotypes exhibit limited cross-protection and transboundary spread potential.

SAT1 differs from other serotypes in that the key antigenic determinants are impacted by changes in the VP1 gene sequence (Dinana et al., 2024). These modifications in the antigenic area directly affect the virus’s capacity to elude the host immune response, which also reduces the efficacy of nonhomologous vaccinations (Jiang et al., 2025). Furthermore, the virus’s adaptation to a variety of host species, including wildlife such as the African buffalo (S. caffer), adds to the complexity of epidemiology and helps maintain the stability of virus circulation in natural reservoirs (Perez-Martin et al., 2022).

However, the absence of equivalent wildlife reservoirs in Indonesia suggests that the dynamics of SAT1 transmission in Southeast Asia may differ from those observed in African ecosystems (Jamal and Belsham, 2013). In Indonesia, if introduction occurs, livestock movement, informal trade, contaminated animal products, and human-mediated transmission would likely become the dominant epidemiological drivers rather than wildlife-mediated maintenance cycles (Rehman et al., 2025). Therefore, African transmission models should be cautiously adapted to local ecological and livestock production conditions (Souley Kouato et al., 2018).

Neutralizing antibodies specific to structural epitopes on the capsid protein play a major role in mediating the protective response to FMDV (Biswal et al., 2020). However, the significant antigenic variety in SAT1 makes it difficult to create a vaccination that is widely effective (Al-Hammadi, 2025). Continuous molecular monitoring is necessary to guarantee the antigenic compatibility of the vaccine because mismatches between vaccine and field strains might lead to protection failure (Han et al., 2023). Vaccine banks should prioritize SAT1 strains representing genetically diverse and antigenically relevant topotypes with demonstrated cross-neutralization potential for emergency preparedness (Wasfy et al., 2025). To maximize protective coverage against potential field strains, selection criteria should include VP1 sequence similarity, phylogenetic clustering, and antigenic matching indicators such as r1-values (Reeve et al., 2010).

Transmission dynamics

There are several intricate ways that FMDV might spread, including interactions between the host, the environment, and human activity (Li et al., 2021). Close contact between infected and susceptible animals is the main way that the virus spreads, especially through respiratory secretions, saliva, vesicular sores, milk, and other excretions that contain a lot of viral particles (Rahman et al., 2025). Animals can release large amounts of virus during the acute phase of infection even before clinical symptoms manifest, raising the possibility of undiscovered transmission (Stenfeldt et al., 2026).

Indirect transmission significantly influences the epidemiology of FMDV in addition to direct transmission (Scharzenberger et al., 2025). The virus can exist in a range of organic materials, including feed, water, soil, and inanimate surfaces (fomites), such as automobiles, workers’ clothing, and livestock equipment (Ryan et al., 2008). Temperature, humidity, and pH all affect the environmental stability of FMDV; cold and humid environments tend to extend viral survival (Mielke and Garabed, 2020). Cross-site transmission is possible even when infected animals are not present (Bessler et al., 2024).

Another significant route is aerosol transmission, especially in situations such as dense populations and favorable wind patterns (Brown et al., 2022). Particularly from animals such as pigs, which are referred to as “amplifier hosts” because of their capacity to produce copious amounts of viral aerosols, virus particles can travel great distances in the air (Dillon, 2011). Cattle, on the other hand, are “indicator hosts” in the early detection of outbreaks because they are more susceptible to infection through inhalation (Gongal et al., 2022).

The local and international mobility of animals and animal products complicates the dynamics of FMDV transmission (Maree et al., 2014). Live livestock trade, the dissemination of poorly processed animal products, and increased human mobility facilitate cross-regional and cross-national spread (Mirzaie et al., 2023). Informal or illicit trade channels frequently present a substantial danger factor in this situation because they are not subject to stringent quarantine and health inspection protocols (Rehman et al., 2025).

Moreover, high-density intensive animal production methods can speed up the pace of transmission within a population (Bellini et al., 2025). The likelihood of virus introduction and amplification at the farm level is increased by inadequate biosecurity measures, such as irregular disinfection and access restriction (Chumsang et al., 2025). Therefore, developing effective control tactics, including risk-based interventions at crucial points in the transmission chain, requires a thorough understanding of FMDV transmission dynamics.

Host range and reservoir

FMDV primarily affects domestic and wild animals with cloven hooves; however, it can also infect a wide variety of hosts (Grubman and Baxt, 2004). The primary livestock animals that are vulnerable to infection include sheep (Ovis aries), goats (Capra hircus), pigs (Sus scrofa domesticus), and cattle (Bos taurus) (Wubshet et al., 2024). Although all species are susceptible to infection, the epidemiological role of each host varies significantly (Humphreys et al., 2025). Pigs act as amplifier hosts because they can produce and shed enormous amounts of virus, especially through aerosols, whereas cattle often operate as indicator hosts because of their high susceptibility to infection (Stenfeldt et al., 2016).

Goats and sheep are examples of small ruminants that frequently show less or even subclinical clinical symptoms, which could act as a covert source of infection (Balinda et al., 2009). This raises the possibility of an unnoticed spread, particularly in conventional livestock systems with little animal health monitoring (Arzt et al., 2018). Additionally, differences in immune responses among species influence the length of virus shedding and infection dynamics (Paton et al., 2018).

Wildlife is essential to the persistence of the virus in the natural world as a reservoir (Hampson and Haydon, 2021). The African buffalo (S. caffer) is one of the primary natural reservoirs for SAT serotypes, particularly SAT1 (Ayebazibwe et al., 2010). This species is a long-term reservoir of the virus because it can sustain a prolonged infection without exhibiting overt clinical symptoms (Humphreys et al., 2025). Interactions between the African buffalo and domestic livestock are crucial for spillover, especially in regions with large livestock systems (Ploquin et al., 2025). Nevertheless, Indonesia does not possess wildlife reservoirs with epidemiological roles comparable to those of African buffalo populations (Triatmojo et al., 2025). Consequently, the absence of these reservoirs may alter the mechanisms sustaining SAT1 circulation, shifting transmission pressure toward livestock-associated pathways rather than wildlife-livestock interfaces (Jori and Etter, 2016).

Although their function as long-term reservoirs is unclear, several other wildlife species, in addition to African buffalo, have also been found to be infected with FMDV (Ayebazibwe et al., 2010). The ability of wildlife to sustain and disseminate the virus is also influenced by ecological factors, including population density, migration patterns, and interspecies interactions (Zhang et al., 2024). Current evidence supporting the long-term maintenance of SAT1 outside African wildlife ecosystems remains limited; therefore, continued genomic surveillance and ecological monitoring are necessary to evaluate whether SAT1 could adapt to alternative transmission systems in Southeast Asia (Mabunda et al., 2026).

Global spread of SAT1: current situation

The global spread of FMDV serotype SAT1 has raised concerns regarding transboundary disease dynamics and their implications for the risk of introduction into disease-free areas, which exhibits a significant shift in epidemiological patterns marked by the expansion of the area of occurrence beyond the historical endemic zone of Sub-Saharan Africa.

Historical distribution

In Sub-Saharan Africa, the FMDV serotype SAT1 has historically maintained a restricted and steady geographic distribution (Sangula et al., 2010). This serotype is mostly associated with savanna environments and continuous encounters between domestic livestock and animals, which facilitate the enzootic cycle of the virus (Jiang et al., 2025). East and Southern African nations, such as Kenya, Tanzania, Zambia, Zimbabwe, Botswana, and South Africa, have long been recognized as important SAT1 endemic regions (Woldemariyam et al., 2023). Ecological conditions, livestock density, and animal transportation all affect regional differences in incidence rates (Aslam and Alkheraije, 2023).

In terms of epidemiology, SAT1 typically persists in wildlife reservoirs, especially African buffalo (S. caffer), which serve as long-term hosts for infection (Jolles et al., 2021). Interactions between domestic livestock and wild reservoirs inside national park borders or conservation zones have maintained an ongoing cycle of virus transmission (Stenfeldt et al., 2025). This pattern leads to seasonal variations in the dynamics of endemic diseases, which are frequently influenced by food availability, animal migration in response to water sources, and climate change (Al-Hammadi, 2025).

Unlike other FMDV serotypes, such as O and A, which are found worldwide, the distribution of SAT1 was comparatively restricted outside of Africa for many years (Lycett et al., 2019). The main causes of this geographic restriction are the existence of particular reservoirs, ecological variables, and restricted live animal trading routes from endemic areas to other regions (Humphreys et al., 2025). However, as global mobility and international livestock trade intensify, this historical stability is starting to shift (Dede et al., 2024).

The latest cross-border spread

In recent years, the distribution of FMDV serotype SAT1 has expanded beyond its usual endemic borders in Sub-Saharan Africa due to changes in epidemiological patterns (Mabunda et al., 2026). With documented occurrences suggesting entry into the Middle East and portions of Western Eurasia, this situation represents a major change in the dynamics of TBDs (Aslam and Alkheraije, 2023). This development is especially concerning because other FMDV serotypes, including O, A, and Asia1, historically predominated in these areas (Hossain et al., 2024).

Antigenic characterization and VP1 gene sequence analysis, which showed phylogenetic proximity to strains circulating in East Africa, were typically used in laboratory confirmation to identify these transboundary outbreaks (Dinana et al., 2026). These findings provide molecular evidence supporting the transboundary dissemination capacity of SAT1 and reinforce concerns regarding its possible spread to non-endemic regions through international trade and livestock mobility networks (Medina and Segundo, 2024). In several epidemic reports, the identification of SAT1 in non-endemic locations was frequently linked to inadequate quarantine system implementation or poorly managed transboundary trade (Byamukama et al., 2025).

The spread of SAT1 outside of Africa has an episodic pattern across time, with outbreaks occurring as intermittent introductions followed by local control measures (Cortey et al., 2019). However, if biosecurity and quick reaction are insufficient, secondary transmission inside impacted areas may occasionally occur, enabling the creation of small transmission chains (Olesen et al., 2020).

This geographical growth also reflects the complexity of logistics systems comprising several international transit hubs and the growing global interconnection of the livestock and animal product trade (An et al., 2025). Additionally, some geopolitical circumstances, such as armed conflict and societal unrest, interfere with animal health surveillance systems, thereby raising the possibility of virus introduction and transmission (Iriarte et al., 2023).

Factors driving the spread

The geographic spread of FMDV serotype SAT1 is influenced by several interrelated factors, such as institutional, anthropogenic, and ecological disease elements (Humphreys et al., 2025). The globalization of trade in live animals and animal products is an important factor because increased cross-border movement raises the risk of virus introduction into areas free of disease (Martínez-López et al., 2008). Complex animal supply chains involving both legal and informal trading, which frequently include transit locations that do not completely enforce biosecurity rules, increase the risk of virus spread (Auty et al., 2019).

Additionally, social unrest and armed conflict play a major role in disrupting disease control systems in many endemic locations (Ahmed et al., 2026). These circumstances may hinder early reporting and outbreak response, impair routine immunization programs, and diminish veterinary surveillance capabilities (Sharan et al., 2023). Therefore, controlling the spread of viruses in animal populations becomes challenging, and they may spread without proper identification (Mashinagu et al., 2024).

The restrictions on the applicability and coverage of immunization programs are another significant concern (Guyver-Fletcher et al., 2025). Antigenic diversity in SAT1 may diminish the efficacy of nonhomologous vaccinations, resulting in an immunological gap in livestock populations (Singh et al., 2019). Virus persistence and transmission are facilitated by mismatches between vaccine and field strains as well as uneven vaccination rates across geographical areas (Mabunda et al., 2026).

However, the identification of epidemics has also been hampered by flaws in epidemiological monitoring systems, such as insufficient laboratory-based early detection and inadequate cross-regional data integration (Iriarte et al., 2023). Accurately tracing the virus’s origins and transmission channels is further hampered by the lack of molecular diagnostic capabilities in certain nations (Longjam et al., 2011).

Risk pathways for their introduction into Indonesia

Given the intricate relationships between animal trade, human mobility, and current biosecurity measures, an analysis of the risk routes for the introduction of the FMDV serotype SAT1 into Indonesia is essential to comprehending the possibility of transboundary disease entry and spread. Table 1 summarizes the various potential risk pathways for the introduction of the FMDV serotype SAT1 into Indonesia, including legal and illegal animal movements, trade and transportation networks, the role of humans as mechanical vectors, live animal markets, and the contribution of the environment and wildlife.

Legal and illegal movements of animals

The transportation of animals across borders, both through official and unofficial trade channels, is a significant contributing element to the introduction and spread of FMDV serotype SAT1 into disease-free regions (Ludi et al., 2016). Importing live livestock, semen, embryos, and animal products through legal routes is governed by health certification and quarantine procedures (Abd El-Rahim et al., 2016). Although these systems are intended to reduce risk, their efficacy is largely dependent on precise detection, compliance with biosecurity regulations, and the alignment of inspection procedures with the current epidemiological dynamics of the nation of origin (van Andel et al., 2021). In infected animals, the virus may evade identification due to pre-certification restrictions or the existence of a subclinical incubation period (Stenfeldt et al., 2025).

Conversely, unauthorized or undocumented routes are more dangerous because they do not go through official veterinary oversight procedures (Blacksell et al., 2023). Uncontrolled land borders for livestock trade, animal product smuggling, and unofficial distribution within local networks can all be significant disease introduction pathways (An et al., 2025). Market demand, economic considerations, and the lax enforcement of regulations in border regions frequently impact these activities (Knight-Jones and Rushton, 2013).

Additionally, regional animal movements, especially commerce between nations in areas with substantial economic integration, increase the complexity of threats (Mabunda et al., 2026). Silent transmission is made possible by interconnected distribution systems, particularly when subclinical-stage animals are transferred before exhibiting obvious clinical symptoms (Scharzenberger et al., 2025). Regional biosecurity systems may have gaps due to disparities in animal health standards between nations (Ahmed et al., 2026).

Trade and transportation networks

The spread of FMDV serotype SAT1 to new areas, particularly Southeast Asia, is greatly increased by international commerce networks and transportation infrastructure (Al-Hammadi, 2025). The movement of goods and animal commodities in large quantities is made possible by regional economic connectedness, especially within trade integration frameworks such as ASEAN (Humphreys et al., 2025). This makes it possible for pathogens to spread through the circulation of goods generated from animals, both those that are well-documented and those that are not (Sutawi et al., 2023).

Given the predominance of international trade via major ports in the Asian region, maritime transportation is an essential part of this dynamic (Aslam and Alkheraije, 2023). Containers, cargo ships, and logistics facilities may act as mechanical vehicles for viral contamination if cleanliness and biosecurity protocols are not followed carefully (Jerab et al., 2026). Because FMDV is stable in some environments, it can survive on inanimate surfaces for a long time to facilitate cross-regional transmission (Brown et al., 2022).

Table 1. Risk pathways for the introduction of the SAT1 serotype of FMDV into Indonesia.

Additionally, on a regional and worldwide level, linked land and air transportation networks speed up the movement of people and the delivery of goods (Tatem et al., 2006). This mobility raises the risk of indirect transmission, especially when it comes to contamination by fomites such as clothing, packaging, or equipment that has come into contact with infectious materials (Auty et al., 2019). In the context of contemporary trade, the intricacy of international logistics networks makes it challenging to identify the source of contamination if a disease is introduced (Tashfeen et al., 2025).

The interconnection of important commercial hubs increases the potential for cross-border spread, particularly when quarantine regulations are inconsistent between nations (Xin et al., 2025). Therefore, if a standardized and integrated biosecurity system is not in place, trade and transportation networks act as both structural and economic drivers that might hasten the transboundary spread of illnesses such as FMDV SAT1 (Mashinagu et al., 2024).

Human-mediated transmission

Humans function as mechanical vectors in the spread of FMDV serotype SAT1 through a variety of actions involving direct or indirect contact with infected animals or contaminated items (Brown et al., 2022). This technique does not include human infection; instead, it occurs when a virus contaminates clothing, cars, equipment, footwear, and body surfaces, which subsequently spreads to vulnerable animal populations (Khairullah et al., 2024).

Those who are actively involved in the livestock production chain, such as farmers, barn workers, veterinarians, and animal transport staff, are more susceptible to transmission (Iriarte et al., 2023). Inadequate human biosecurity practices, such as cleaning shoes and wearing protective clothing, can make it easier for viruses to spread between farms (Chumsang et al., 2025). Additionally, one significant route for indirect transmission at the local level is the usage of shared equipment without proper sanitation (Brown et al., 2022).

Regional and global human mobility also increases the risk of virus spread (Lessani et al., 2024). If hygiene precautions are not carefully followed, interregional travel that includes stops at livestock facilities or animal markets may become crucial locations for the introduction of pathogens (Rehman et al., 2025). Employees who handle the logistics of animals and animal products are particularly vulnerable as mechanical carriers in the international trade context, particularly if they come from endemic regions (Longjam et al., 2011).

Furthermore, because of the high concentration of animals from different origins and the frequent interactions between people and animals, activities in live animal marketplaces greatly increase the accumulation of risk (Kedang et al., 2025). These circumstances foster cross-contamination, particularly if regular disinfection and hygiene practices are not practiced (Chumsang et al., 2025).

Environmental and wildlife trails

Although trade and domestic animal mobility play a larger role in the introduction and survival of FMDV serotype SAT1, environmental and wildlife channels still play a significant role in transboundary disease risk assessments (Sangula et al., 2010). Indirect infection through soil, water, feed, or surfaces exposed to secretions from infected animals is possible because the virus may endure specific environmental conditions for a brief amount of time, especially at low temperatures and high humidity (Scharzenberger et al., 2025). However, exposure to ultraviolet radiation, temperature changes, and microbiological activity tends to have a major impact on the stability of FMDV in the open environment, all of which lower viral vitality (Kamolsiripichaiporn et al., 2007).

Natural reservoirs are crucial to FMDV ecology in wildlife, especially in endemic regions of Africa where African buffalo (S. caffer) act as long-term hosts for infection (van Schalkwyk et al., 2016). However, no known wildlife reservoirs with a comparable epidemiological significance for SAT1 exist in places like Indonesia (Jamal and Belsham, 2013). However, the possible role of local wild species as spillover hosts must still be considered, especially in areas that separate significant livestock systems from natural ecosystems (Stenfeldt et al., 2025).

Indirect contact between domestic animals and wildlife might occur in pastures, water sources, or conservation areas adjacent to livestock activities (Mashinagu et al., 2024). Although very little empirical data support the role of non-endemic species in sustaining the FMDV SAT1 cycle, these conditions could theoretically form an ecological bridge for disease transmission (Abousenna, 2026).

Indonesia’s vulnerability and preparedness measures

A thorough analysis of Indonesia’s susceptibility to the possible introduction of FMDV serotype SAT1 is necessary, considering the country’s livestock system dynamics, the volume of animal trade, and differences in surveillance and biosecurity capabilities, which together affect the degree of readiness for transboundary disease threats.

The current status of FMD in Indonesia

Following an epidemic in 2022 that signaled the disease’s reappearance after a protracted period of clinical freedom, FMD has once again become a major problem in Indonesia (Salman et al., 2025). Since then, FMDV has been detected in several provinces, with serotype O predominating, indicating that it can quickly spread in domestic livestock systems, especially in dairy and beef cattle populations (Hartaputera et al., 2024). The virus continues to circulate in small regions with low to moderate endemicity despite intensive control measures (Woldemariyam et al., 2023).

Mass vaccination, animal movement limitations, and heightened monitoring of livestock transit between regions are examples of control measures implemented (Singh et al., 2019). Although vaccination coverage, prompt delivery, and the vaccine strain’s compatibility for the field virus all affect its efficacy, emergency vaccination is a crucial step in lowering the incidence of clinical cases (Orsel and Bouma, 2009). Additionally, active surveillance and a strengthened reporting system have been put in place to speed up the identification of new cases and more precisely map the distribution of the disease (Ahmed et al., 2026).

Biosecurity implementation at the farm level still exhibits notable regional diversity in terms of non-vaccine management, especially in relation to resource capacity and farmer compliance. This contributes to local transmission persistence (Fountain et al., 2023). Although issues with unofficial distribution channels remain, livestock movement surveillance and animal quarantine procedures have been strengthened in the interim (Xin et al., 2025).

Immunity and the vaccination gap

Antigenic incompatibilities between the field virus and available vaccination strains are the main factor influencing the susceptibility of livestock populations to FMDV serotype SAT1 (Ludi et al., 2016). Vaccination campaigns in Indonesia have mostly concentrated on locally prevalent serotypes, especially serotype O, which has resulted in little cross-protection against SAT1 (Kurniawan et al., 2024). Because important antigenic determinants, especially the VP1 capsid protein, differ significantly among FMDV serotypes, there is little to no immunological cross-reactivity (Zewdie et al., 2023). This limitation highlights the importance of establishing vaccine banks containing antigenically characterized SAT1 strains with broad cross-reactivity profiles (Mahapatra and Parida, 2018). To reduce the risk of vaccine failure during emergency outbreaks, antigen selection should be continuously updated using genomic surveillance, VP1 sequence analysis, phylogenetic mapping, and in vitro antigenic matching studies (Elrashedy et al., 2025a).

Regional variations in vaccination coverage have also been linked to the immunity gap in livestock populations (Knight-Jones et al., 2016). Variations in vaccination rates, inconsistent booster intervals, and limited vaccine distribution in some regions have led to the formation of susceptible pockets that could potentially become starting points for amplification if a new virus is introduced (Belsham, 2020). In addition, species, physiological state, and previous antigen exposure can all affect an individual’s immune response (Medina et al., 2018).

Post-vaccination immunity against FMDV is short-lived and necessitates recurring boosts to keep neutralizing antibody titers at protective levels (Sitt et al., 2019). However, the paucity of information regarding the antigenic match between the vaccine candidates employed and the SAT1 strains that are circulating worldwide is a significant obstacle to vaccination preparation planning (Zewdie et al., 2023).

Diagnostic and surveillance capacity

Surveillance systems and diagnostic capabilities are essential for the early detection and reaction to the possible arrival of FMDV serotype SAT1 (Longjam et al., 2011). Currently, Indonesia’s diagnostic capabilities include both molecular techniques, such as reverse transcription polymerase chain reaction, for quick virus identification and conventional methods, such as enzyme-linked immunosorbent assay-based antigen detection and antibody detection, for serological confirmation (Dinana et al., 2023). Nevertheless, only a few reference laboratories use full genomics-based methods, such as whole-genome sequencing and real-time phylogenetic analysis (Abd El Rahman et al., 2020).

These restrictions affect the speed and precision of detecting viral genetic alterations, which are essential for tracking the possible emergence of novel serotypes, such as SAT1 (Al-Hammadi, 2025). In the absence of sufficient genomic data, the capacity to map transmission channels and perform molecular tracing is subpar (Klein, 2009). Additionally, there is still a delay between case discovery and the use of control measures due to incomplete real-time coordination of laboratory data integration with field surveillance systems (Hamza et al., 2025).

From the standpoint of epidemiological surveillance, passive reporting continues to predominate in the current system, with unevenly distributed active monitoring across regions providing assistance (Arjkumpa et al., 2020). The sensitivity of the system in early infection detection is further influenced by differences in laboratory infrastructure, human resource capacity, and access to contemporary diagnostic technology (Siswani et al., 2025).

Strategic elements for enhancing readiness for the launch of FMDV SAT1 include bolstering genomic-based diagnostic capability, growing the national laboratory network, and incorporating digital surveillance data (Dinana et al., 2026). Additionally, this strategy makes it possible to implement a more responsive early warning system based on molecular data, which facilitates quicker decision-making in the management of outbreaks (Abousenna, 2026).

Biosecurity practices

Implementation of biosecurity procedures is a key element in stopping the introduction and spread of FMDV serotype SAT1 at the farm level and at points of entry (Singh et al., 2019). Biosecurity at the farm level includes a range of preventive measures intended to reduce contact between susceptible animals and infectious agents, such as managing livestock entry and exit, controlling vehicle and human access, maintaining facility sanitation, and isolating new animals prior to their introduction into the main population (Arjmand et al., 2025). The degree of technology use, farm management capabilities, and adherence to standard operating procedures have a significant impact on the effectiveness of these indicators (Syihabuddin et al., 2025).

The implementation of biosecurity in small- to medium-sized livestock systems is frequently adaptive and varies, impacted by a lack of resources, expertise, and supporting infrastructure (Mehmedi et al., 2025). Therefore, different industrial units have different levels of protection, which leads to risk heterogeneity within a regional population (Otieno et al., 2023). On the other hand, biosecurity is more structured in intensive production systems, albeit maintaining uniform application across all operational lines remains difficult (Chumsang et al., 2025).

Animal quarantine systems, which act as a major filter against the possible transboundary entry of diseases, are used to conduct biosecurity at the border level (Mielke et al., 2023). Verification of animal health records, clinical examinations, laboratory testing, and setting up observation periods in compliance with relevant rules are among the measures implemented (Garner et al., 2021). However, the sensitivity of diagnostic techniques, the capacity for early detection, and the ability to detect infections in the subclinical stage all play a significant role in the efficacy of this system (Stenfeldt et al., 2025).

Beyond technological issues, the biggest obstacles to biosecurity implementation are uniform regulation enforcement and cross-administrative region integration (Mashinagu et al., 2024). Gaps in coordination and the presence of unofficial trade routes may make control less effective at crucial junctures (Stenfeldt and Arzt, 2020). Therefore, bolstering risk-based biosecurity and harmonizing operational standards across the animal distribution chain are key components in lowering the possibility of FMDV SAT1 entering the country’s livestock system (Mabunda et al., 2026).

Prevention and control strategies

To successfully break the transmission chain and lower the risk of disease introduction and dissemination, efforts to prevent and control the FMDV serotype SAT1 require an integrated approach that incorporates surveillance-based interventions, vaccination, and biosecurity.

Strengthening the surveillance system

A key component of the plan to stop and manage the spread of FMDV serotype SAT1 is bolstering the surveillance system, mainly through improved early detection capabilities and real-time virus dynamics monitoring (Sangula et al., 2010). In addition to identifying clinical cases, an efficient surveillance system incorporates a risk-based strategy that combines laboratory, epidemiological, and animal mobility data into a thorough analytical framework (Humphreys et al., 2025).

Combining passive and active surveillance can improve early detection (Gunasekara et al., 2021). Active monitoring entails methodical field inspections in high-risk communities, such as border regions, livestock trading hubs, and regions with dense animal populations, whereas passive surveillance relies on case reports from farmers and animal health professionals (Arjkumpa et al., 2020). Combining these two methods can enhance the system’s sensitivity to early infection event detection (Adamchick et al., 2021).

Genomic surveillance is essential to combat the threat posed by novel serotypes, such as SAT1 (Ahmed et al., 2026). Precise molecular characterization of the virus, including determination of its origin, mode of introduction, and genetic alterations that may impact virulence and antigenicity, is made possible by whole-genome sequencing and phylogenetic analysis (Sulistyaningrum et al., 2024). These genomic data also aid in the assessment of vaccination suitability by tracking antigenic drift and lineage replacement (Upadhyaya et al., 2021).

Additionally, integrating digital-based information technologies to speed up data flow from the field level to analysis centers is necessary to boost contemporary surveillance (Zakharova et al., 2021). Interoperability between animal health organizations and real-time reporting systems can enhance the speed and coordination of outbreak responses (Oltean et al., 2025). In this regard, the integration of data on human factors, the environment, and animal health into a single integrated monitoring system is another application of the One Health method (Basheer et al., 2025).

Vaccination strategy

A vaccine strategy is essential for controlling FMDV, especially considering the possible introduction of the highly variable SAT1 serotype (Paton et al., 2009). Because of insufficient cross-protection between FMDV serotypes, vaccination efficacy is primarily based on the degree of antigenic match between vaccine strains and field viruses (Dekker et al., 2020). Consequently, a strain-matching methodology is required to develop an immunization strategy that adjusts to global epidemiological dynamics (Sahoo et al., 2025).

Continuous molecular monitoring data, such as VP1 sequence analysis and phylogenetic mapping to identify circulating viral clusters, are necessary to adapt the vaccine composition to the SAT1 lineage (Elrashedy et al., 2025b). Candidate vaccine strains with the best antigenic match to field isolates were identified using these data (Hassan et al., 2022). A more accurate vaccine selection process can be facilitated using an antigenic mapping approach to depict antigenic associations across strains (Bari et al., 2014).

In addition to vaccine production, the readiness plan calls for the creation and upkeep of an emergency vaccination bank (Lu et al., 2022). This technique speeds up the mass immunization response by enabling quick vaccine availability in the case of an introduction or outbreak hazard (Hamza et al., 2025). The availability of a representative array of antigens against several FMDV lineages, including SAT1, and a well-organized delivery mechanism are critical to vaccine banks’ efficacy (Singh et al., 2019).

Additionally, enhancing the cold chain logistics system, ranking target populations according to epidemiological risk, and routinely assessing the degree of population immunity by serological surveillance are all necessary for implementing an ideal vaccination strategy (Park et al., 2021). An integrated approach is essential to guarantee that vaccination treatments are not just reactive but also preventive and grounded in current scientific data (Al-Hammadi, 2025).

Enhanced biosecurity

The main preventive strategy for reducing the danger of the introduction and spread of the FMDV serotype SAT1 is enhanced biosecurity, which breaks the chain of transmission at crucial junctures in animal production and distribution networks (Abousenna, 2026). A risk-based biosecurity strategy that is adapted to the features of the livestock system, animal migration patterns, and the degree of regional vulnerability is necessary for effective biosecurity implementation (González-Gordon et al., 2023).

Stricter access controls for people, cars, and equipment, as well as uniform cleaning and disinfection practices, are all part of enhanced biosecurity at the farm level (Chumsang et al., 2025). One of the most important steps in preventing the introduction of clinically undiagnosed infectious agents is isolating new animals (quarantine holding) before integrating them into the main population (Rehman et al., 2025). Internal livestock movement management must also be controlled to prevent animal groups with disparate health conditions from coming into contact with one another EFSA Panel on Animal Health and Welfare (AHAW) et al., 2021.

Strengthening border quarantine systems is the main line of defense against the transboundary entrance of diseases at the regional and national levels (Rehman et al., 2025). The optimization of quarantine functions involves enhancing laboratory diagnostic skills to identify subclinical diseases, validating animal health records, and expanding clinical inspection capacity (Longjam et al., 2011). In high-risk scenarios, rapid diagnostic technologies and digital tracking systems can increase detection effectiveness and speed up decision-making (Rios et al., 2018).

The degree of compliance and uniformity of application throughout the cattle value chain also influence the success of biosecurity (Renault et al., 2021). Epidemiological gaps that raise the risk of virus introduction might result from gaps in the application of operational standards, especially in small-scale manufacturing systems and unofficial trading routes (Humphreys et al., 2025). Therefore, enhancing cross-sectoral monitoring, human resource capacity building, and regulatory harmonization are required to develop a sustainable and adaptable biosecurity system against the threat of FMDV SAT1 (Maehira and Spencer, 2019).

Preparedness and response planning

The main objectives of preparedness and response planning, which is a strategic component in reducing the danger of FMDV serotype SAT1 introduction and dissemination, are to speed up identification, restrict transmission, and lessen epidemiological and economic effects (Saeed et al., 2015). A scenario-based readiness approach that considers several potential routes of introduction, transmission dynamics, and response capabilities at the local and national levels is the foundation of an efficient preparedness system (Bellini et al., 2025).

An outbreak simulation is an essential technique for evaluating a system’s readiness for exceptional occurrences (Roche et al., 2015). These tasks include field reaction drills, epidemiological modeling, and agency coordination assessment (An et al., 2025). Simulations can be used to identify potential operational challenges, such as case identification delays, a lack of logistical resources, and communication gaps between response units (Chen and Meng, 2025). The outcomes of these assessments serve as a foundation for improving handling procedures and boosting the effectiveness of the response system (Al-Hammadi, 2025).

A rapid response system is necessary to provide timely interventions after the first case discovery (Dinana et al., 2023). The availability of emergency supplies, such as vaccinations and disinfectants, a trained fast response team, a real-time reporting system, and standardized outbreak control protocols are important elements of this system (Meckawy et al., 2022). In particular, for diseases such as FMD that have high transmission characteristics, the response time is critical in preventing the initial spread (Stenfeldt et al., 2026).

Supporting evidence-based decision-making requires the integration of diagnostic laboratory networks with epidemiological information systems (González-Gordon et al., 2023). The use of digital platforms for data reporting and analysis makes early trend detection and quicker reaction protocol activation possible (Kashem et al., 2025). The ability of preparedness systems to adapt to changing transmission patterns and antigenic ambiguity is essential considering the threat posed by novel serotypes, such as SAT1 (Pomeroy et al., 2015).

One Health and regional collaboration

The One Health method is an integrative paradigm that unifies ecosystems, human health, and animal health into a single, cohesive system of analysis and intervention (Guo et al., 2023). This method is applicable to FMDV serotype SAT1 because the intricate relationships between livestock systems, human mobility, animal trade, and environmental factors influence the dynamics of disease dissemination in addition to the biology of the virus (Danasekaran, 2024). Cross-sectoral integration makes more thorough risk identification possible, especially at the interface between local inhabitants and the larger ecosystem (Eltahir et al., 2024).

Public health organizations, environmental agencies, and animal health authorities must work closely together to exchange data, harmonize surveillance techniques, and respond to disease outbreaks (Gerilovych et al., 2026). This strategy also improves early detection capabilities by integrating epidemiological data from many sectors, enabling quicker identification and better coordinated control of possible introductions of novel serotypes such as SAT1 (Elrashedy et al., 2025c).

Given that FMD is a transboundary disease that crosses national administrative borders, regional cooperation is essential (Rahman et al., 2025). Coordinating policies to regulate the transportation of animals and animal products, standardizing diagnostic procedures, and exchanging epidemic data are examples of cooperation within the ASEAN framework and other international networks (Blacksell et al., 2019). This approach also entails fortifying a collaborative reporting system and a genetic information-sharing platform to facilitate real-time tracking of virus evolution (Makau et al., 2025a).

Additionally, regional cooperation facilitates the development of member nations’ capacities through technology transfer, human resource training, and technical assistance in creating monitoring and epidemic response systems (Mphande-Nyasulu et al., 2024). This synergy is becoming increasingly crucial to combat the threat posed by viruses with substantial antigenic variation, such as SAT1, which call for a coordinated strategy to close capacity gaps between regions (Mahapatra et al., 2016).

Policy implications and recommendations

Strengthening an adaptable and risk-based animal health governance system is necessary to address the policy consequences of the possible introduction of FMDV serotype SAT1 (Al-Hammadi, 2025). Harmonizing national biosecurity laws, which include standardizing quarantine practices, monitoring animal movements, and enforcing compliance throughout the livestock distribution chain, is the top priority (Arjmand et al., 2025). Cross-sectoral coordination structures must be in place for this strategy to be implemented consistently at the national and regional levels (Al-Hammadi, 2025).

Research and development spending is essential, especially in genomic epidemiology, molecular virology, and antigenic matching-based vaccine development (Klein, 2009). To improve the early detection and characterization of potentially introduced virus strains, the reference laboratory infrastructure, including genome sequencing and bioinformatics analysis capacity, must be strengthened (Humphreys et al., 2025).

To increase international cooperation, it is necessary to actively participate in regional and global surveillance networks, such as exchanging genetic data, reporting outbreaks in real time, and working together to develop and distribute vaccines (Mabunda et al., 2026). This cooperation is essential to improve early warning systems against the threat of highly mobile transboundary diseases (Paton et al., 2009).

Beyond technical considerations, a plan for communicating risks to stakeholders is essential to the effectiveness of control measures EFSA Panel on Animal Health and Welfare (AHAW) et al., 2021 Compliance with biosecurity procedures and immunization programs can be increased by providing accurate, consistent, and scientifically based information to livestock producers, industry participants, and the general public (Rehman et al., 2025). Therefore, an integrated policy strategy that includes legislation, scientific capacity, international cooperation, and risk communication is the fundamental basis for reducing the FMDV SAT1 threat (Mabunda et al., 2026).

Future perspectives

According to the epidemiological dynamics of FMDV, the SAT1 serotype is not the only concern; other SAT serotypes (SAT2 and SAT3), which have similar antigenic evolution patterns, may also appear or reemerge (Jiang et al., 2025). The genetic diversity of the SAT group increases the likelihood of lineage alterations, which could affect the efficacy of vaccines and change the geographic distribution of the disease (Li et al., 2023). Therefore, it is essential to continuously study molecular evolution to predict changes in virus dynamics worldwide (Aslam and Alkheraije, 2023).

Artificial intelligence and machine learning technologies are becoming increasingly important to improve FMD surveillance systems (Makau et al, 2025b). This technology makes large-scale epidemiological data analysis possible, making it possible to find risk patterns, forecasting disease development, and spotting outbreak abnormalities faster than traditional techniques (González-Gordon et al., 2023). The accuracy of early warning systems may be increased by including environmental variables, cattle migration, and genomic data in prediction models (Wang et al., 2021).

In addition to biological and technological aspects, global climate change is predicted to affect the dynamics of FMDV dissemination through variations in temperature, humidity, and mechanical vector distribution (Dede et al., 2024). While shifting seasonal patterns may affect livestock movements and population interactions, warmer and more humid environmental conditions may boost the short-term stability of viruses outside of their hosts (Mielke and Garabed, 2020). Indirect effects, such as modifications to cattle production systems brought on by climate stress, may also increase vulnerability to the introduction and spread of illness (Stenfeldt et al., 2025).

---

Conclusion

The FMDV serotype SAT1 poses a potential threat to endemic-free countries because of its high transmission capacity, antigenic variability, and history of cross-border expansion from endemic areas in Africa. Although direct evidence of sustained SAT1 circulation in Southeast Asia remains limited, molecular surveillance data and reports of transboundary outbreaks indicate that SAT serotypes possess the capacity for geographic expansion beyond their historical endemic zones. The likelihood of the virus spreading to areas with sizable livestock populations and diverse livestock systems, such as Indonesia, is increased by the complexity of introduction pathways, which include animal trade, human mobility, and international logistics networks.

Strengthening genomic-based surveillance, modifying vaccination plans according to antigenic compatibility, and establishing uniform biosecurity across the production and distribution chain are all necessary for effective preparedness. Future preparedness strategies should prioritize region-specific transmission modeling and the development of vaccine banks containing antigenically matched SAT1 strains to improve the rapid response capability in the event of introduction.

---

Acknowledgment

The authors gratefully acknowledge the financial support provided by the Indonesia Endowment Fund for Education (LPDP) through its collaborative research funding program and the National Research and Innovation Agency (BRIN), Indonesia, for providing institutional support and research facilities.

Funding

Indonesia Endowment Fund for Education (LPDP) through its collaborative research funding program and the National Research and Innovation Agency (BRIN), Indonesia.

Author’s contributions

HN, ARK, NLPID, AG, and RSA drafted the manuscript. AW, SMN, FE, RAA, and DN revised and edited the manuscript. MS, IS, MKJK, and IF prepared and critically checked this manuscript. SW, RS, EO, and NSJL have edited the references. All authors have read and approved the final version of the manuscript.

Conflict of interest

The authors have no potential conflicts of interest to declare.

Data availability

The data used in this study were obtained from publicly available sources on the internet. All relevant data are cited within the article, and the corresponding author can provide additional details upon reasonable request.

---

References

Abd El Rahman, S., Hoffmann, B., Karam, R., El-Beskawy, M., Hamed, M.F., Forth, L.F., Höper, D. and Eschbaumer, M. 2020. Sequence analysis of Egyptian foot-and-mouth disease virus field and vaccine strains: intertypic recombination and evidence for accidental release of virulent virus. Viruses 12(9), 990.

Abd El-rahim, I.H., Asghar, A.H., Mohamed, A.M. and Fat’hi, S.M. 2016. The impact of importation of live ruminants on the epizootiology of foot and mouth disease in Saudi Arabia. Rev. Sci. Tech. 35(3), 769–778.

Abousenna. 2026. Current situation and emerging foot-and-mouth disease virus lineages in Egypt: historicment SAT1 Introduction, and Vaccine Policy Implications. Vet. Immunol. Immunopathol. 292(1), 111063.

Adamchick, J., Rich, K.M. and Perez, A.M. 2021. Assessment of the risk of foot and mouth disease among beef cattle at slaughter from east african production systems. Viruses 13(12), 2407.

Adnyana, I.M., Utomo, B., Eljatin, D.S. and Sudaryati, N.L. 2023. One Health approach and zoonotic diseases in Indonesia: urgency of implementation and challenges. Narra J. 3(3), 257.

Ahmed, M.J., Hossain, M.I., Rahman, M.A., Bhuiyan, M.I.H., Bhandari, P., Alam, K.E., Chalise, R., Kaderi, I.J., Rahi, M.A.I., Tarin, T., Alam, M.J., Adesola, R.O. and Hossain, D. 2026. Prevalence and serotype distribution of foot and mouth disease (FMD) Virus in Asian Countries: a systematic review and meta-analysis. Vet. Med. Int. 1(1), 5492291.

Al-Hammadi, M.A. 2025. Current trends and challenges in the management of foot and mouth disease in Saudi Arabia: a review. Open Vet. J. 15(5), 1907–1933.

An, Q., Lv, Y., Li, Y., Sun, Z., Gao, X. and Wang, H. 2025. Global foot-and-mouth disease risk assessment based on multiple spatial analysis and ecological niche model. Vet. Q. 45(1), 1–11.

Arjkumpa, O., Yano, T., Prakotcheo, R., Sansamur, C. and Punyapornwithaya, V. 2020. Epidemiology and national surveillance system for foot and mouth disease in cattle in Thailand during 2008–2019. Vet. Sci. 7(3), 99.

Arjmand, A., Bani-Yaghoub, M., Corkran, K., Pandit, P.S. and Aly, S.S. 2025. Assessing the impact of biosecurity compliance on farmworker and livestock health within a one health modeling framework. One Health 20(1), 101023.

Arzt, J., Belsham, G.J., Lohse, L., Bøtner, A. and Stenfeldt, C. 2018. Transmission of foot-and-mouth disease from persistently infected carrier cattle to naive cattle via transfer of oropharyngeal fluid. mSphere 3(5), 365.

Aslam, M. and Alkheraije, K.A. 2023. The prevalence of foot-and-mouth disease in Asia. Front. Vet. Sci. 10(1), 1201578.

Auty, H., Mellor, D., Gunn, G. and Boden, L.A. 2019. The risk of foot and mouth disease transmission posed by public access to the countryside during an outbreak. Front. Vet. Sci. 6(1), 381.

Ayebazibwe, C., Mwiine, F.N., Tjørnehøj, K., Balinda, S.N., Muwanika, V.B., Ademun Okurut, A.R., Belsham, G.J., Normann, P., Siegismund, H.R. and Alexandersen, S. 2010. The role of African buffalos (Syncerus caffer) in the maintenance of foot-and-mouth disease in Uganda. BMC. Vet. Res. 6(1), 54.

Balinda, S.N., Tjørnehøj, K., Muwanika, V.B., Sangula, A.K., Mwiine, F.N., Ayebazibwe, C., Masembe, C., Siegismund, H.R. and Alexandersen, S. 2009. Prevalence estimates of antibodies towards foot-and-mouth disease virus in small ruminants in Uganda. Transbound. Emerg. Dis. 56(9-10), 362–371.

Bari, F.D., Parida, S., Tekleghiorghis, T., Dekker, A., Sangula, A., Reeve, R., Haydon, D.T., Paton, D.J. and Mahapatra, M. 2014. Genetic and antigenic characterisation of serotype A FMD viruses from East Africa to select new vaccine strains. Vaccine 32(44), 5794–5800.

Basheer, A., Tran, M., Khan, B., Jentner, W., Wendelboe, A., Vogel, J., Kuhn, K., Wimberly, M.C. and Ebert, D. 2025. Comprehensive review of One Health systems for emerging infectious disease detection and management. One Health 21(1), 101253.

Bellini, S., Scaburri, A., Tironi, M., Cappa, V., Mannelli, A. and Alborali, G.L. 2025. Simulating the spread of foot-and-mouth disease in densely populated areas as part of contingency plans to establish the best control options. Pathogens 14(9), 933.

Belsham, G.J. 2020. Towards improvements in foot-and-mouth disease vaccine performance. Acta. Vet. Scand. 62(1), 20.

Bessler, A.L., Nayee, S., Garabed, R., Krug, P., Obrycki, J. and Rodriguez, L. 2024. Surviving the summer: foot-and-mouth disease virus survival in U.S. regional soil types at high ambient temperatures. Front. Vet. Sci. 11(1), 1429760.

Biswal, J.K., Subramaniam, S., Ranjan, R., Vanderwaal, K., Sanyal, A., Pattnaik, B. and Singh, R.K. 2020. Differential antibody responses to the major antigenic sites of FMD virus serotype O after primo-vaccination, multiply-vaccination and after natural exposure. Infect. Genet. Evol. 78(1), 104105.

Blacksell, S.D., Dhawan, S., Kusumoto, M., Le, K.K., Summermatter, K., O’Keefe, J., Kozlovac, J., Almuhairi, S.S., Sendow, I., Scheel, C.M., Ahumibe, A., Masuku, Z.M., Bennett, A.M., Kojima, K., Harper, D.R. and Hamilton, K. 2023. The biosafety research road map: the search for evidence to support practices in the laboratory-foot and mouth disease virus. Appl. Biosafety. 28(4), 199–215.

Blacksell, S.D., Siengsanan-Lamont, J., Kamolsiripichaiporn, S., Gleeson, L.J. and Windsor, P.A. 2019. A history of FMD research and control programmes in Southeast Asia: lessons from the past informing the future. Epidemiol. Infect. 147(1), 171.

Brown, E., Nelson, N., Gubbins, S. and Colenutt, C. 2022. Airborne transmission of foot-and-mouth disease virus: a review of past and present perspectives. Viruses 14(5), 1009.

Byamukama, B., Amin, A., Mwiine, F.N. and Ekiri, A.B. 2025. Epidemiology and control strategies for foot-and-mouth disease in livestock and wildlife in Uganda: systematic review. Vet. Res. Commun. 49(4), 227.

Chen, L. and Meng, Q.H. 2025. Advancing laboratory diagnostics for future pandemics: challenges and innovations. Pathogens 14(11), 1135.

Chumsang, P., Singhla, T. and Chaisowwong, W. 2025. Qualitative risk assessment of foot-and-mouth disease virus introduction and transmission to dairy farms via raw milk transportation in thailand: a scenario-based approach. Vet. Sci. 12(7), 623.

Cortey, M., Ferretti, L., Pérez-Martín, E., Zhang, F., De Klerk-lorist, L.M., Scott, K., Freimanis, G., Seago, J., Ribeca, P., Van Schalkwyk, L., Juleff, N.D., Maree, F.F. and Charleston, B. 2019. Persistent infection of African buffalo (Syncerus caffer) with foot-and-mouth disease virus: limited viral evolution and no evidence of antibody neutralization escape. J. Virol. 93(15), 563.

Danasekaran, R. 2024. One health: a holistic approach to tackling global health issues. Indian. J. Community. Med. 49(2), 260–263.

Dede, M., Sunardi, S., Widiawaty, M.A., Ismail, A., Lam, K.C., Afriana, N.N., Susilowati, S., Husodo, T., Sukriah, E. and Susiati, H. 2024. Spatial distribution and environmental factors analysis of foot-and-mouth disease (FMD) in West Java, Indonesia. Cogent Food Agric. 11(1), 2440549.

Dekker, A., Sanz-Bernardo, B., Singanallur, N.B., Ludi, A.B., Horsington, J., Eblé, P.L., King, D.P. and Vosloo, W. 2020. Cross-protection induced by a A/MAY/97 emergency vaccine against intra-serotype heterologous challenge with a foot-and-mouth disease virus from the A/ASIA/G-VII lineage. Vaccines. 8(1), 24.

Dillon, M.B. 2011. Skin as a potential source of infectious foot and mouth disease aerosols. Proc. Biol. Sci. 278(1713), 1761–1769.

Dinana, Z., Rahmahani, J., Suwarno., Hamid, I.S., Al-Arif, M.A., Yunus, M., Tyasningsih, W., Ihsan, I.S., Maharani, A.T., Maulana, F.K., Said, N.S., Fahrodi, D.U. and Rantam, F.A. 2026. Molecular evolution and epidemiological dynamics of foot-and-mouth disease virus O/ME-SA/Ind-2001e Circulating in East Java, Indonesia, in 2022-2025. Vet. Med. Int. 1(1), 6526830.

Dinana, Z., Rantam, F.A., Suwarno, S., Mustofa, I., Rahmahani, J. and Kusnoto, K. 2023. Detection of foot and mouth disease virus in cattle in Lamongan and Surabaya, Indonesia using RT-PCR method. J. Med. Vet. 6(2), 191–196.

Dinana, Z., Suwarno, S., Mustofa, I., Rahmahani, J., Kusnoto, K., Maharani, A.T., Fitria, A.L., Witaningrum, A.M., Maulana, F.K., Said, N.S., Fahrodi, D.U. and Rantam, F.A. 2024. Molecular characterization of VP1 gene during the foot-and-mouth disease virus outbreak in East Java, Indonesia, in 2022. Vet. World. 17(11), 2469–2476.

Duchatel, F., Bronsvoort, B.M.D.C. and Lycett, S. 2019. Phylogeographic analysis and identification of factors impacting the diffusion of foot-and-mouth disease virus in Africa. Front. Ecol. Evol. 7(1), 371.

EFSA Panel on Animal Health and Welfare (AHAW), Nielsen, S.S., Alvarez, J., Bicout, D.J., Calistri, P., Canali, E., Drewe, J.A., Garin-Bastuji, B., Rojas, J.L.G., Schmidt, C.G., Herskin, M., Michel, V., Chueca, M.A.M., Padalino, B., Pasquali, P., Sihvonen, L.H., Spoolder, H., Ståhl, K., Velarde, A., Viltrop, A., Winckler, C., Clercq, K.D., Gubbins, S., Klement, E., Stegeman, J.A., Antoniou, S.E., Aznar, I., Broglia, A., Papanikolaou, A., der Stede, Y.V., Zancanaro, G. and Roberts, H.C. 2021. Scientific Opinion on the assessment of the control measures for category A diseases of Animal Health Law: Foot and Mouth Disease. EFSA J. 19(6), e06632.

Ekanayaka, P., Shin, S.H., Weeratunga, P., Lee, H., Kim, T.H., Chathuranga, K., Subasinghe, A., Park, J.H. and Lee, J.S. 2021. Foot-and-mouth disease Virus 3C Protease Antagonizes Interferon Signaling and C142T Substitution Attenuates the FMD Virus. Front. Microbiol. 12(1), 737031.

Elrashedy, A., Mousa, W., Nayel, M., Salama, A., Zaghawa, A., Elsify, A. and Hasan, M.E. 2025. Systematic review and meta-analysis of the effectiveness of polypeptide, virus-like particles, and viral vector vaccines for foot-and-mouth disease (2020-2025). Sci. Rep. 15(1), 39370.

Elrashedy, A., Nayel, M., Salama, A., Zaghawa, A., Badr, A. and Hasan, M.E. 2025b. Phylogenetic and evolutionary analysis of VP1 coding sequences of foot-and-mouth disease virus serotypes A, O, and SAT2 in Egypt. Virol. J. 23(1), 24.

Elrashedy, A., Nayel, M., Salama, A., Zaghawa, A., El-Shabasy, R.M. and Hasan, M.E. 2025a. Foot-and-mouth disease: genomic and proteomic structure, antigenic sites, serotype relationships, immune evasion, recent vaccine development strategies, and future perspectives. Vet. Res. 56(1), 78.

Eltahir, Y.M., Ishag, H.Z.A., Wadsworth, J., Hicks, H.M., Knowles, N.J., Mioulet, V., King, D.P., Mohamed, M.S., Bensalah, O.K., Yusof, M.F., Gasim, E.F.M., Hammadi, Z.M.A., Shah, A.A.M., Abdelmagid, Y.A., Gahlan, M.A.M.E., Kassim, M.F., Kayaf, K., Zahran, A. and Nuaimat, M.M.A. 2024. Molecular epidemiology of foot-and-mouth disease viruses in the Emirate of Abu Dhabi, United Arab Emirates. Vet. Sci. 11(1), 32.

Fountain, J., Manyweathers, J., Brookes, V.J. and Hernandez-Jover, M. 2023. Understanding biosecurity behaviors of Australian beef cattle farmers using the ten basic human values framework. Front. Vet. Sci. 10(1), 1072929.

Gao, Y., Sun, S.Q. and Guo, H.C. 2016. Biological function of Foot-and-mouth disease virus non-structural proteins and non-coding elements. Virol. J. 13(1), 107.

Garner, G., Vosloo, W., Tapsuwan, S., Bradhurst, R., Seitzinger, A.H., Breed, A.C. and Capon, T. 2021. Comparing surveillance approaches to support regaining free status after a foot-and-mouth disease outbreak. Prev. Vet. Med. 194(1), 105441.

Gerilovych, A., Shevchenko, N., Pishchanskyi, O., Aliekseieva, H., Rosada, M., Gerilovych, I. and Okaievych, O. 2026. Operationalizing the One Health approach in a conflict-affected setting: a scientometric review of policy foundations, systemic gaps, and future pathways in Ukraine. Vet. World 19(1), 389–408.

Gizaw, D., Senbata, B., Fentie, A., Bilata, T., Negessu, D., Muluneh, A., Shegu, D., Ashenafi, H., Knowles, N.J., Wadsworth, J., Mioulet, V., Hicks, H.M., Legesse, M., Kassa, T. and King, D.P. 2025. Serotype diversity and molecular characterization of foot-and-mouth disease viruses from outbreaks in Ethiopia (2019-2023): re-Emergence of SAT 2 After 30 Years. Transbound. Emerg. Dis. 1(1), 6670343.

Gongal, G., Rahman, H., Thakuri, K.C. and Vijayalakshmy, K. 2022. An Overview of Transboundary Animal Diseases of Viral Origin in South Asia: what Needs to Be Done?. Vet. Sci. 9(11), 586.

González-Gordon, L., Porphyre, T., Muwonge, A., Nantima, N., Ademun, R., Ochwo, S., Mwiine, N.F., Boden, L., Muhanguzi, D. and Bronsvoort, B.M.D.C. 2023. Identifying target areas for risk-based surveillance and control of transboundary animal diseases: a seasonal analysis of slaughter and live-trade cattle movements in Uganda. Sci. Rep. 13(1), 18619.

Grubman, M.J. and Baxt, B. 2004. Foot-and-mouth disease. Clin. Microbiol. Rev. 17(2), 465–493.

Gunasekara, U., Bertram, M.R., Dung, D.H., Hoang, B.H., Phuong, N.T., Hung, V.V., Long, N.V., Minh, P.Q., Vu, L.T., Dong, P.V., Perez, A., Vanderwaal, K. and Arzt, J. 2021. Use of slaughterhouses as sentinel points for genomic surveillance of foot-and-mouth disease virus in Southern Vietnam. Viruses 13(11), 2203.

Guo, Z.Y., Zheng, J., Li, S.Z. and Zhou, X.N. 2023. Orientation of One Health development: think globally and act locally. Sci. One. Health. 2(1), 100042.

Guyver-Fletcher, G., Gorsich, E.E., Jewell, C. and Tildesley, M.J. 2025. Controlling endemic foot-and-mouth disease: vaccination is more important than movement bans. A simulation study in the Republic of Turkey. Infect. Dis. Model. 10(2), 702–715.

Hampson, K. and Haydon, D. 2021. Persistent pathogens and wildlife reservoirs. Science 374(6563), 35–36.

Hamza, I.A., Mao, K., Gao, C., Hamza, H. and Zhang, H. 2025. Elements of viral outbreak preparedness: lessons, strategies, and future directions. Viruses 18(1), 50.

Han, J.H., Yoo, D.S. and Lee, C.M. 2023. Effect of a mismatched vaccine against the outbreak of a novel FMD strain in a pig population. Animals (Basel). 13(19), 3082.

Hartaputera, I.N.S.T., Suryadarma, I.G.A., Mahardika, I.G.N.K. and Suardana, I.W. 2024. Serotype of foot-and-mouth disease virus in cattle circulated in Asia region including Indonesia: a literature review. Indones. Med. Vet. 13(3), 296–312.

Hassan, A., Aboezz, Z., El-Habbaa, A., Shahein, M., Hagag, N. and Sharawy, S. 2022. Phylogenetic analysis of VP1 foot and mouth disease virus strains circulating between 2018-2020, Egypt. Benha. Vet. Med. J. 42(2), 7–13.

Hossain, A., Alam, K.M.M., Akter, S., Hossain, M.A. and Sultana, M. 2024. A comprehensive and single-use foot-and-mouth disease sero-surveillance prototype employing rationally designed multiple viral antigens. Sci. Rep. 14(1), 25236.

Humphreys, J.M., Stenfeldt, C., King, D.P., Knight-Jones, T., Perez, A.M., Vanderwaal, K., Sanderson, M.W., Di Nardo, A., Jemberu, W.T., Pamornchainavakul, N. and Arzt, J. 2025. Epidemiology and economics of foot-and-mouth disease: current understanding and knowledge gaps. Vet. Res. 56(1), 141.

Iriarte, M.V., Gonzáles, J.L., De Freitas Costa, E., Gil, A.D. and De Jong, M.C.M. 2023. Main factors associated with foot-and-mouth disease virus infection during the 2001 FMD epidemic in Uruguay. Front. Vet. Sci. 10(1), 1070188.

Jamal, S.M. and Belsham, G.J. 2013. Foot-and-mouth disease: past, present and future. Vet. Res. 44(1), 116.

Jerab, J.G., Biebaut, E., Berge, A.C., Chantziaras, I. and Dewulf, J. 2026. Are Visitor and Personnel Downtime Restrictions an Effective Biosecurity Measure to Prevent the Indirect Transmission of Pathogens to Livestock?. Animals (Basel). 16(2), 205.

Jiang, S., Yang, S., Zhang, X., Fang, Y., Guo, Z., Ma, Z., Wei, X., Guo, K., El-Ansary, R.E., Bayasgalan, C., Wubshet, A.K. and Zhang, J. 2025. Evolutionary and structural insights into VP1 epitopes of representative SAT-type FMDV strains: implications for candidate vaccine selection. Vet. Res. 56(1), 227.

Jolles, A., Gorsich, E., Gubbins, S., Beechler, B., Buss, P., Juleff, N., De Klerk-lorist, L.M., Maree, F., Perez-Martin, E., Van Schalkwyk, O.L., Scott, K., Zhang, F., Medlock, J. and Charleston, B. 2021. Endemic persistence of a highly contagious pathogen: foot-and-mouth disease in its wildlife host. Science 374(6563), 104–108.

Jori, F. and Etter, E. 2016. Transmission of foot and mouth disease at the wildlife/livestock interface of the Kruger National Park, South Africa: can the risk be mitigated?. Prev. Vet. Med. 126(1), 19–29.

Kamolsiripichaiporn, S., Subharat, S., Udon, R., Thongtha, P. and Nuanualsuwan, S. 2007. Thermal inactivation of foot-and-mouth disease viruses in suspension. Appl. Environ. Microbiol. 73(22), 7177–7184.

Kashem, M.A., Ambagala, T., Hole, K., Yang, M., Nfon, C. and Babiuk, S. 2025. A Simplified mAb-Based Antigen Detection Assay for Rapid Serotyping of Foot-and-Mouth Disease Virus. Viruses 17(6), 761.

Kedang, V.M.K., Permatasari, I., Chanchaidechachai, T. and Inchaisri, C. 2025. Spatial-temporal distribution and risk factors of foot-and-mouth disease outbreaks in Java Island, Indonesia from 2022 to 2023. BMC. Vet. Res. 21(1), 180.

Khairullah, A.R., Kurniawan, S.C., Effendi, M.H., Silaen, O.S.M., Moses5, I.B., Hasib, A., Ramandinianto, S.C., Afnani, D.A., Widodo, A., Riwu, K.H.P., Az Zahra, R.L. and Yanestria, S.M. 2024. The danger of foot and mouth disease in livestock – a review. Borneo. J. Resour. Sci. Technol. 14(2), 173–187.

Klein, J. 2009. Understanding the molecular epidemiology of foot-and-mouth-disease virus. Infect. Genet. Evol. 9(2), 153–161.

Knight-Jones, T.J.D. and Rushton, J. 2013. The economic impacts of foot and mouth disease - what are they, how big are they and where do they occur?. Prev. Vet. Med. 112(3–4), 161–173.

Knight-Jones, T.J.D., Gubbins, S., Bulut, A.N., Stärk, K.D.C., Pfeiffer, D.U., Sumption, K.J. and Paton, D.J. 2016. Mass vaccination, immunity and coverage: modelling population protection against foot-and-mouth disease in Turkish cattle. Scientific Rep. 6(1), 22121.

Kurniawan, Y., Tyasningsih, W., Rahmahani, J., Puspitasari, Y., Kusnoto, K., Azzahra, F., Tobing, T.M., Aswin, A., Diyantoro, D., Maulana, F.K., Susilowati, H., Kuncorojakti, S. and Rantam, F.A. 2024. Protein characterization of an Indonesian isolate of foot and mouth disease virus inactivated with formaldehyde and binary ethylenimine. Vet. World 17(8), 1836–1845.

Lessani, M.N., Li, Z., Jing, F., Qiao, S., Zhang, J., Olatosi, B. and Li, X. 2024. Human mobility and the infectious disease transmission: a systematic review. Geo-spat. Inf. Sci. 27(6), 1824–1851.

Li, C., Shi, J., Wang, H., Rivera-Serrano, E.E., Yang, D., Zhou, G., Sun, C., Cameron, C.E. and Yu, L. 2020. Polymerase Fidelity Contributes to Foot-and-Mouth Disease Virus Pathogenicity and Transmissibility In Vivo. J. Virol. 95(1), 15–69.

Li, K., Wang, C., Yang, F., Cao, W., Zhu, Z. and Zheng, H. 2021. Virus-Host Interactions in Foot-and-Mouth Disease Virus Infection. Front. Immunol. 12(1), 571509.

Li, Q., Wubshet, A.K., Wang, Y., Heath, L. and Zhang, J. 2023. B and T cell epitopes of the incursionary foot-and-mouth disease virus serotype SAT2 for vaccine development. Viruses 15(3), 797.

Longjam, N., Deb, R., Sarmah, A.K., Tayo, T., Awachat, V.B. and Saxena, V.K. 2011. A Brief Review on diagnosis of foot-and-mouth disease of livestock: conventional to molecular tools. Vet. Med. Int. 1(1), 905768.

Lu, Z., Yu, S., Wang, W., Chen, W., Wang, X., Wu, K., Li, X., Fan, S., Ding, H., Yi, L. and Chen, J. 2022. Development of foot-and-mouth disease vaccines in recent years. Vaccines (Basel). 10(11), 1817.

Ludi, A., Ahmed, Z., Pomeroy, L.W., Pauszek, S.J., Smoliga, G.R., Moritz, M., Dickmu, S., Abdoulkadiri, S., Arzt, J., Garabed, R. and Rodriguez, L.L. 2016. Serotype diversity of foot-and-mouth-disease virus in livestock without history of vaccination in the far north region of cameroon. Transbound. Emerg. Dis. 63(1), e27–e38.

Lycett, S., Tanya, V.N., Hall, M., King, D.P., Mazeri, S., Mioulet, V., Knowles, N.J., Wadsworth, J., Bachanek-Bankowska, K., Ngu Ngwa, V., Morgan, K.L. and Bronsvoort, B.M.D.C. 2019. The evolution and phylodynamics of serotype A and SAT2 foot-and-mouth disease viruses in endemic regions of Africa. Sci. Rep. 9(1), 5614.

Mabunda, G.P., Selaledi, L. and Nemukondeni, N. 2026. Global synthesis of serotype diversity, transmission dynamics and socioeconomic impacts of foot-and-mouth disease. Discov. Virus. 3(1), 4.

Maehira, Y. and Spencer, R.C. 2019. Harmonization of biosafety and biosecurity standards for high-containment facilities in low- and middle-income countries: an approach from the perspective of occupational safety and health. Front. Public Health 7(1), 249.

Mahapatra, M. and Parida, S. 2018. Foot and mouth disease vaccine strain selection: current approaches and future perspectives. Expert Rev. Vaccines 17(7), 577–591.

Mahapatra, M., Statham, B., Li, Y., Hammond, J., Paton, D. and Parida, S. 2016. Emergence of antigenic variants within serotype A FMDV in the Middle East with antigenically critical amino acid substitutions. Vaccine 34(27), 3199–3206.

Makau, D.N., Arzt, J. and Vanderwaal, K. 2025a. Opportunities for machine learning to predict cross-neutralization in FMDV serotype O. PLos Comput. Biol. 21(9), e1013491.

Makau, D.N., Arzt, J. and Vanderwaal, K. 2025b. Tracing the spread and phylogeography of foot-and-mouth disease virus across East and the Horn of Africa. Virus. Evol. 11(1), 73.

Maree, F.F., Kasanga, C.J., Scott, K.A., Opperman, P.A., Melanie, C., Sangula, A.K., Raphael, S., Yona, S., Wambura, P.N., King, D.P., Paton, D.J. and Rweyemamu, M.M. 2014. Challenges and prospects for the control of foot-and-mouth disease: an African perspective. Vet. Med. (Auckl). 5(1), 119–138.

Martínez-López, B., Perez, A.M., De La Torre, A. and Rodriguez, J.M.S.V. 2008. Quantitative risk assessment of foot-and-mouth disease introduction into Spain via importation of live animals. Prev. Vet. Med. 86(1–2), 43–56.

Mashinagu, M.M., Wambura, P.N., King, D.P., Paton, D.J., Maree, F., Kimera, S.I., Rweyemamu, M.M. and Kasanga, C.J. 2024. Challenges of controlling foot-and-mouth disease in pastoral settings in Africa. Transbound. Emerg. Dis. 1(1), 2700985.

Maxim, L., Mazzocchi, M., Van Den Broucke, S., Zollo, F., Robinson, T., Rogers, C., Vrbos, D., Zamariola, G. and Smith, A. 2021. Technical assistance in the field of risk communication. EFSA. J. 19(4), 6574.

Meckawy, R., Stuckler, D., Mehta, A., Al-Ahdal, T. and Doebbeling, B.N. 2022. Effectiveness of early warning systems in the detection of infectious diseases outbreaks: a systematic review. BMC. Public. Health. 22(1), 2216.

Medina, G.N. and Segundo, F.D.S. 2024. Virulence and immune evasion strategies of FMDV: implications for vaccine design. Vaccines 12(9), 1071.

Medina, G.N., Segundo, F.D.S., Stenfeldt, C., Arzt, J. and De Los Santos, T. 2018. The Different tactics of foot-and-mouth disease virus to evade innate immunity. Front. Microbiol. 9(1), 2644.

Mehmedi, B., Iatrou, A.M., Yildiz, R., Lamont, K., Da Costa, M.R., De Nardi, M., Allepuz, A., Niine, T., Niemi, J.K. and Saegerman, C. 2025. Economic perspectives on farm biosecurity: stakeholder challenges and livestock species considerations. Agriculture 15(21), 2288.

Mielke, S.R. and Garabed, R. 2020. Environmental persistence of foot-and-mouth disease virus applied to endemic regions. Transbound. Emerg. Dis. 67(2), 543–554.

Mielke, S.R., Lendzele, S., Delgado, A.H., Abdoulmoumini, M., Dickmu, S. and Garabed, R. 2023. Patterns of foot-and-mouth disease virus detection in environmental samples in an endemic setting. Front. Vet. Sci. 10(1), 1157538.

Mirzaie, K., Mowlaei, S., Arsevska, E., Ahmadi, B.V., Ambrosini, F., Rosso, F. and Chevanne, E. 2023. Analysis of livestock mobility and implications for the risk of foot-and-mouth disease virus spread in Iran. Ruminants 3(4), 299–323.

Mohamadin, M., Manzoor, R., Elolimy, A., Abdelmegeid, M., Mosad, S. and Abd El Rahman, S. 2025. Advancements in antiviral approaches against foot-and-mouth disease virus: a comprehensive review. Front. Vet. Sci. 12(1), 1574193.

Mphande-Nyasulu, F.A., Yap, N.J., Teo, C.H., Chang, L.Y. and Tay, S.T. 2024. Outbreak preparedness and response strategies in ASEAN member states: a scoping review. IJID Reg. 12(1), 100430.

Nielsen, S.S., Alvarez, J., Bicout, D.J., Calistri, P., Canali, E., Drewe, J.A., Garin-Bastuji, B., Gonzales Rojas, J.L., Gortázar Schmidt, C., Herskin, M., Michel, V., Miranda Chueca, M.A., Padalino, B., Pasquali, P., Sihvonen, L.H., Spoolder, H., Ståhl, K., Velarde, A., Viltrop, A., Winckler, C., De Clercq, K., Gubbins, S., Klement, E., Stegeman, J.A., Antoniou, S.E., Aznar, I., Broglia, A., Papanikolaou, A., Van Der Stede, Y., Zancanaro, G. and Roberts, H.C. 2021. Scientific opinion on the assessment of the control measures for category A diseases of Animal Health Law: foot and mouth disease. EFSA. J. 19(6), 6632.

Olesen, A.S., Belsham, G.J., Bruun Rasmussen, T., Lohse, L., Bødker, R., Halasa, T., Boklund, A. and Bøtner, A. 2020. Potential routes for indirect transmission of African swine fever virus into domestic pig herds. Transbound. Emerg. Dis. 67(4), 1472–1484.

Oltean, H.N., Lipton, B., Black, A., Snekvik, K., Haman, K., Buswell, M., Baines, A.E., Rabinowitz, P.M., Russell, S.L., Shadomy, S., Ghai, R.R., Rekant, S., Lindquist, S. and Baseman, J.G. 2025. Developing a one health data integration framework focused on real-time pathogen surveillance and applied genomic epidemiology. One Health Outlook 7(1), 9.

Orsel, K. and Bouma, A. 2009. The effect of foot-and-mouth disease (FMD) vaccination on virus transmission and the significance for the field. Can. Vet. J. 50(10), 1059–1063.

Otieno, W.A., Nyikal, R.A., Mbogoh, S.G. and Rao, E.J.O. 2023. Adoption of farm biosecurity practices among smallholder poultry farmers in Kenya - An application of latent class analysis with a multinomial logistic regression. Prev. Vet. Med. 217(1), 105967.

Park, M.Y., Han, Y.J., Choi, E.J., Kim, H., Pervin, R., Shin, W., Kwon, D., Kim, J.M. and Pyo, H.M. 2021. Post-vaccination monitoring to assess foot-and-mouth disease immunity at population level in Korea. Front. Vet. Sci. 8(1), 673820.

Paton, D.J., Gubbins, S. and King, D.P. 2018. Understanding the transmission of foot-and-mouth disease virus at different scales. Curr. Opin. Virol. 28(1), 85–91.

Paton, D.J., Sumption, K.J. and Charleston, B. 2009. Options for control of foot-and-mouth disease: knowledge, capability and policy. Philos. Trans. R. Soc. Lond. B. Biol. Sci. 364(1530), 2657–2667.

Perez-Martin, E., Beechler, B., Zhang, F., Scott, K., De Klerk-lorist, L.M., Limon, G., Dugovich, B., Gubbins, S., Botha, A., Hetem, R., Van Schalkwyk, L., Juleff, N., Maree, F.F., Jolles, A. and Charleston, B. 2022. Viral dynamics and immune responses to foot-and-mouth disease virus in African buffalo (Syncerus caffer). Vet. Res. 53(1), 63.

Ploquin, O., Grosbois, V., Ndlovu, M., Ndozore, S., Munzamba, M., Porovha, E., Nkomo, K., Basso, O., Corbel, G., Shumba, R., Mhlanga, M.D., Mwandirigana, E., Musekiwa, B., Takayindisa, E., Loisier, A., Fritz, H., Liégeois, F., Caron, A., Prugnolle, F. and Miguel, E. 2025. Foot-and-mouth disease dynamics in multi-species livestock systems at the interface of African protected areas. Vet. Res. 56(1), 58.

Pomeroy, L.W., Bjørnstad, O.N., Kim, H., Jumbo, S.D., Abdoulkadiri, S. and Garabed, R. 2015. Serotype-Specific transmission and waning immunity of endemic foot-and-mouth disease virus in Cameroon. PLos One. 10(9), 136642.

Rahman, M.A., Zereen, F., Rana, M.L., Hossain, M.G., Shimada, M. and Saha, S. 2025. Foot-and-mouth disease in Asia. Virus. Res. 351(1), 199514.

Reeve, R., Blignaut, B., Esterhuysen, J.J., Opperman, P., Matthews, L., Fry, E.E., De Beer, T.A.P., Theron, J., Rieder, E., Vosloo, W., O’Neill, H.G., Haydon, D.T. and Maree, F.F. 2010. Sequence-based prediction for vaccine strain selection and identification of antigenic variability in foot-and-mouth disease virus. PLos Comput. Biol. 6(12), e1001027.

Rehman, S., Ullah, S., Abuzahra, M., Effendi, M.H., Budiastuti, B., Kholik, K., Munawarah, M., Zaman, A., Rahman, A.U., Malik, M.I., Ullah, S. and Rustam, S.A. 2025. Determination of risk factors for foot and mouth disease emergence in East Java, Indonesia. Open Vet. J. 15(5), 2049–2058.

Renault, V., Humblet, M.F., Pham, P. and Saegerman, C. 2021. Biosecurity at Cattle Farms: strengths, Weaknesses, Opportunities and Threats. Pathogens 10(10), 1315.

Rios, L., Perera, C.L., Coronado, L., Relova, D., Álvarez, A.M., Ganges, L., Díaz De Arce, H., Núñez, J.I. and Pérez, L.J. 2018. Multi-target strategy for pan/foot-and-mouth disease virus (FMDV) detection: a combination of sequences analysis, in Silico predictions and laboratory diagnostic evaluation. Front. Vet. Sci. 5(1), 160.

Roche, S.E., Garner, M.G., Sanson, R.L., Cook, C., Birch, C., Backer, J.A., Dubé, C., Patyk, K.A., Stevenson, M.A., Yu, Z.D., Rawdon, T.G. and Gauntlett, F. 2015. Evaluating vaccination strategies to control foot-and-mouth disease: a model comparison study. Epidemiol&. Infection 143(6), 1256–1275.

Ryan, E., Mackay, D. and Donaldson, A. 2008. Foot-and-mouth disease virus concentrations in products of animal origin. Transbound. Emerg. Dis. 55(2), 89–98.

Saeed, A., Kanwal, S., Arshad, M., Ali, M., Shaikh, R.S. and Abubakar, M. 2015. Foot-and-mouth disease: overview of motives of disease spread and efficacy of available vaccines. J. Anim. Sci. Technol. 57(1), 10.

Sahoo, S., Lee, H.K. and Shin, D. 2025. An integrated structural and immunoinformatic approach to design a multi-epitope based vaccine against the foot-and-mouth disease virus. Sci. Rep. 15(1), 35911.

Salman, A., Susetya, H., Indarjulianto, S. and Budiyanto, A. 2025. Spatiotemporal analysis of the re-emerging foot and mouth disease outbreak in Central Java, Indonesia. Open Vet. J. 15(6), 2703–2714.

Sangula, A.K., Belsham, G.J., Muwanika, V.B., Heller, R., Balinda, S.N., Masembe, C. and Siegismund, H.R. 2010. Evolutionary analysis of foot-and-mouth disease virus serotype SAT 1 isolates from east Africa suggests two independent introductions from southern Africa. BMC. Evol. Biol. 10(1), 371.

Scharzenberger, S., Humphreys, J.M., Rigney, C., Freifeld, A., Stenfeldt, C. and Arzt, J. 2025. Epidemiologic consequences of preclinical transmission of foot-and-mouth disease virus in cattle. Front. Vet. Sci. 12(1), 1651091.

Sharan, M., Vijay, D., Yadav, J.P., Bedi, J.S. and Dhaka, P. 2023. Surveillance and response strategies for zoonotic diseases: a comprehensive review. Sci. One Health 2(1), 100050.

Singh, R.K., Sharma, G.K., Mahajan, S., Dhama, K., Basagoudanavar, S.H., Hosamani, M., Sreenivasa, B.P., Chaicumpa, W., Gupta, V.K. and Sanyal, A. 2019. Foot-and-mouth disease virus: immunobiology, advances in vaccines and vaccination strategies addressing vaccine failures-an Indian perspective. Vaccines (Basel). 7(3), 90.

Siswani, S., Hatta, M., Muflihanah, M., Muhiddin, N.M., Ekawasti, F., Damayanti, R., Ahmad, R.Z., Intan, P.R., Fitriana, F., Desem, M.I. and Rinendyaputri, R. 2025. Sensitivity of specimen type for diagnosing foot-and-mouth diseases in cattle using one-step real-time polymerase chain reaction (3D gene). Open Vet. J. 15(1), 252–260.

Sitt, T., Kenney, M., Barrera, J., Pandya, M., Eckstrom, K., Warner, M., Pacheco, J.M., Larocco, M., Palarea-Albaladejo, J., Brake, D., Rieder, E., Arzt, J., Barlow, J.W. and Golde, W.T. 2019. Duration of protection and humoral immunity induced by an adenovirus-vectored subunit vaccine for foot-and-mouth disease (FMD) in Holstein steers. Vaccine 37(42), 6221–6231.

Souley Kouato, B., De Clercq, K., Abatih, E., Dal Pozzo, F., King, D.P., Thys, E., Marichatou, H. and Saegerman, C. 2018. Review of epidemiological risk models for foot-and-mouth disease: implications for prevention strategies with a focus on Africa. PLos One. 13(12), 208296.

Stenfeldt, C. and Arzt, J. 2020. The carrier conundrum; a review of recent advances and persistent gaps regarding the carrier state of foot-and-mouth disease virus. Pathogens 9(3), 167.

Stenfeldt, C., Diaz-san Segundo, F., De Los Santos, T., Rodriguez, L.L. and Arzt, J. 2016. The Pathogenesis of Foot-and-Mouth Disease in Pigs. Front. Vet. Sci. 3(1), 41.

Stenfeldt, C., Eschbaumer, M., Humphreys, J., Medina, G.N. and Arzt, J. 2025. The pathogenesis of foot-and-mouth disease virus: current understandings and knowledge gaps. Vet. Res. 56(1), 119.

Stenfeldt, C., Humphreys, J.M. and Arzt, J. 2026. Incubation phase transmission of foot-and-mouth disease virus in cattle: experimental evidence and simulated impacts. Sci. Rep. 16(1), 4023.

Sulistyaningrum, E., Wibawa, H. and Wibowo, M.H. 2024. Identification and molecular characterization of foot and mouth disease virus based on VP1 gene fragments in Madura cattle and Ongole grade cattle. Trop. Anim. Sci. J. 47(2), 170–179.

Sutawi, S., Wahyudi, A., Malik, A., Suyatno, S., Hidayati, A., Rahayu, I.D. and Hartatie, E.S. 2023. Re-emergence of foot and mouth disease outbreak in Indonesia: a review. Adv. Anim. Vet. Sci. 11(2), 264–271.

Syihabuddin, M.Y., Andarwati, S., Guntoro, B. and Putra, A.R.S. 2025. Evaluating foot and mouth disease vaccination services through assessment of beef cattle farmers’ satisfaction in Sleman Regency. Trop. Anim. Sci. J. 48(1), 83–92.

Tashfeen, A., Rahman, M. and Gul, A.S.T. 2025. Clinical diagnosis, epidemiological patterns, and prophylactic strategies for foot-and-mouth disease in ruminants and swine. J. Health Wellness Community Res. 3(14), 984.

Tatem, A.J., Rogers, D.J. and Hay, S.I. 2006. Global transport networks and infectious disease spread. Adv. Parasitology 62(1), 293–343.

Triatmojo, A., Guntoro, B., Strausz, P., Muzayyanah, M.A.U., Agustiar, R. and Kusza, S. 2025. Socioeconomic impact of foot and mouth disease outbreaks on smallholder cattle farmers in Yogyakarta, Indonesia. Vet. Sci. 12(6), 542.

Upadhyaya, S., Mahapatra, M., Mioulet, V. and Parida, S. 2021. Molecular basis of antigenic drift in serotype o foot-and-mouth disease viruses (2013-2018) from Southeast Asia. Viruses 13(9), 1886.

Van Andel, M., Tildesley, M.J. and Gates, M.C. 2021. Challenges and opportunities for using national animal datasets to support foot-and-mouth disease control. Transbound. Emerg. Dis. 68(4), 1800–1813.

Van Schalkwyk, O.L., Knobel, D.L., De Clercq, E.M., De Pus, C., Hendrickx, G. and Van Den Bossche, P. 2016. Description of events where african buffaloes (Syncerus caffer) strayed from the endemic foot-and-mouth disease zone in South Africa, 1998-2008. Transbound. Emerg. Dis. 63(3), 333–347.

Vosloo, W., Boshoff, K., Dwarka, R. and Bastos, A. 2002. The possible role that buffalo played in the recent outbreaks of foot-and-mouth disease in South Africa. Ann. N. Y. Acad. Sci. 969(1), 187–190.

Wang, J., Chen, J., Zhang, S., Ding, Y., Wang, M., Zhang, H., Liang, R., Chen, Q. and Niu, B. 2021. Risk assessment and integrated surveillance of foot-and-mouth disease outbreaks in Russia based on Monte Carlo simulation. BMC Vet. Res. 17(1), 268.

Wasfy, M., Bazid, A.H., Nayel, M., Ata, E.B., Elfeil, W.K., Attia, M. and Elsayed, M. 2025. Immunogenicity of a foot-and-mouth disease (FMD) vaccine against serotypes O, A, SAT-2, and Asia-1 in the Middle East and many parts of Africa, Southeast Asia and Europe. Virol. J. 22(1), 98.

Woldemariyam, F.T., Kariuki, C.K., Kamau, J., De Vleeschauwer, A., De Clercq, K., Lefebvre, D.J. and Paeshuyse, J. 2023. Epidemiological dynamics of foot-and-mouth disease in the horn of Africa: the role of virus diversity and animal movement. Viruses 15(4), 969.

Wubshet, A.K., Werid, G.M., Teklue, T., Zhou, L., Bayasgalan, C., Tserendorj, A., Liu, J., Heath, L., Sun, Y., Ding, Y., Wang, W., Zaberezhny, A.D., Liu, Y. and Zhang, J. 2024. Foot and mouth disease vaccine efficacy in Africa: a systematic review and meta-analysis. Front. Vet. Sci. 11(1), 1360256.

Xin, J., Lan, S., Ai, J., Zeng, B., Xin, A., Ye, L., Zuo, W., Li, Y. and Han, D. 2025. Risk assessment and prevention of foot-and-mouth disease transmission from laos to China. Vet. Sci. 12(2), 92.

Zakharova, O.I., Korennoy, F.I., Iashin, I.V., Toropova, N.N., Gogin, A.E., Kolbasov, D.V., Surkova, G.V., Malkhazova, S.M. and Blokhin, A.A. 2021. Ecological and socio-economic determinants of livestock animal leptospirosis in the Russian Arctic. Front. Vet. Sci. 8(1), 65867.

Zewdie, G., Akalu, M., Tolossa, W., Belay, H., Deresse, G., Zekarias, M. and Tesfaye, Y. 2023. A review of foot-and-mouth disease in Ethiopia: epidemiological aspects, economic implications, and control strategies. Virol. J. 20(1), 299.

Zhang, S., Chai, R., Hu, Y., Joka, F.R., Wu, X., Wang, H. and Wang, X. 2024. Unveiling the spatial distribution and transboundary pathways of FMD serotype O in Western China and its bordering countries. PLos One. 19(8), 306746.