Phenotypic characteristics of adult swamp buffalo in smallholder systems of Central Lombok, Indonesia

Muhammad Muhsinin¹, Maskur¹, Tapaul Rozi¹, Milda Tantrini² and Muhammad Salman Al Farizi³

¹Laboratory of Animal Breeding and Genetics, Faculty of Animal Science, University of Mataram, Mataram 83125, Indonesia
muhsinin@unram.ac.id
²Undergraduate Student, Faculty of Animal Science, University of Mataram, Mataram 83125, Indonesia
³Postgraduate Student, Department of Animal Production and Technology, Faculty of Animal Science, IPB University, Bogor 16680, Indonesia

Abstract

Swamp buffalo are central to smallholder mixed farming systems in Central Lombok, Indonesia, where they provide draught power, meat, manure and savings for rural households, yet systematic phenotypic information on these locally adapted animals is scarce. This study aimed to describe the qualitative and quantitative phenotypic characteristics of adult swamp buffalo kept by smallholder farmers and to derive simple descriptors that can be used in on-farm management, selection and conservation. Sixty adult buffalo were selected using purposive sampling based on age and health status and were grouped by sex and age class (36–48 and 48–60 months). Qualitative traits (head shape and colour, horn shape, neck type, neck collar line, body hair, skin, leg and tail hair colour) were recorded by visual observation and quantitative traits (shoulder height, body length, chest circumference, hip height, horn length, tail length and bodyweight) were measured using standard morphometric procedures and field weight estimation methods. Frequencies and percentages were calculated for qualitative traits and means, standard deviations and coefficients of variation for quantitative traits by sex and age group. Adult swamp buffalo showed a dominant qualitative profile characterised by oval heads, backward-curved horns, grey body hair, black skin, white legs and black-brown tail hair. Older animals (48–60 months) had larger body dimensions and higher bodyweight than younger adults and males were larger than females, reflecting typical sexual dimorphism. Coefficients of variation for main body measurements were mostly below 15 percent, indicating relatively homogeneous body size, while horn and tail length were more variable. Bodyweight increased with chest circumference, confirming that simple tape measurements can be used to estimate weight under field conditions. The phenotypic baseline generated in this study supports the use of simple descriptors and body measurements for on-farm selection, weight estimation and conservation planning of swamp buffalo in tropical smallholder mixed farming systems.

Key words: body measurements, indigenous buffalo, phenotypic traits, smallholder farmers, mixed farming systems, tropical livestock

Introduction

Swamp buffalo (Bubalus bubalis carabanensis) are important livestock in many parts of Asia, including Indonesia, where they provide draught power, meat, manure and socio-cultural value to rural communities. In Central Lombok, West Nusa Tenggara, buffalo are typically kept by smallholder farmers under extensive and semi-intensive systems and contribute to household livelihoods as well as to the conservation of local animal genetic resources. They are integrated into mixed crop–livestock farming, where they use locally available forages and crop residues and return nutrients to the soil through manure, while also functioning as a form of savings and risk management for smallholder households.

Phenotype reflects the expression of an animal's genotype under specific environmental conditions and includes qualitative traits such as coat colour, horn shape, head shape and body markings and quantitative traits such as linear body measurements and bodyweight. Qualitative traits are useful as visual descriptors for distinguishing and describing populations while quantitative traits are more closely related to production potential, functional adaptation and overall conformation. Phenotypic characterization using both types of traits is widely recognised as a first step in the management, conservation and improvement of animal genetic resources in smallholder farming systems (Bousbia et al 2021; Deribe et al 2021).

Recent work on Indonesian buffalo has shown variation in body size and shape between regions and identified chest girth, body length and height at withers as important traits for describing morphometric diversity (Nafiu et al 2025; Saputra and Anggraeni 2023). Studies on indigenous buffalo in Southeast Asia report medium to large body size, grey to dark coat colours and compact conformation and emphasise the value of phenotypic descriptors for local breed management and performance evaluation (Maulana et al 2023; Prihandini et al 2023; Suhardi et al 2022). A recent review of Indonesian indigenous buffalo biodiversity highlighted that phenotypic uniformity within regions can be high but genetic differentiation among regions is often substantial and stressed the need for region-specific phenotypic baselines to support conservation programmes (Prihandini et al 2023). Genetic studies have also revealed that maternal genetic diversity within swamp buffalo populations may be limited due to founder effects and limited gene flow, which underscores the importance of phenotypic monitoring for conservation planning (Brocklebank et al 2025). In addition, studies on Indonesian buffalo have shown that simple linear measurements such as chest girth and body length can be used to predict bodyweight and can therefore support on-farm selection and management in low-input systems (Wari et al 2025).

Information on the phenotypic characteristics of adult swamp buffalo in Central Lombok, especially regarding the distribution of qualitative traits and the variability of quantitative measurements within the local population, is still scarce. This gap makes it difficult to compare populations across regions, to monitor changes over time and to design locally adapted breeding or conservation strategies that are consistent with sustainable smallholder farming (Prihandini et al 2023; Suhardi et al 2022). Generating a clear phenotypic baseline for adult swamp buffalo in Central Lombok can therefore support practical selection, bodyweight estimation without scales and informed decision-making for conservation and sustainable use in low-input systems.

The aim of the present study was to characterize the qualitative and quantitative phenotypic traits of adult swamp buffalo kept by smallholder farmers in Central Lombok, Indonesia. It was hypothesised that the population would show a dominant phenotypic profile with relatively low variability in major body dimensions but with some variation in traits such as horn and tail length that could be relevant for selection and conservation in smallholder systems.

Materials and methods

Study area and farming systems

The study was conducted in several sub‑districts of Central Lombok Regency, West Nusa Tenggara Province, Indonesia, including areas where swamp buffalo are commonly integrated into mixed crop–livestock systems. Central Lombok lies in the central part of Lombok Island and is characterised by a tropical monsoon climate with a distinct rainy season from approximately November to March and a dry season from April to October. The study sites are located at low to medium altitudes, where smallholder farmers cultivate rice and maize and keep swamp buffalo under extensive grazing and tethering systems on communal and private pastureland. A map showing the location of Central Lombok within West Nusa Tenggara Province and the approximate position of the sampled sub‑districts has been included (Figure 1) to assist readers who are not familiar with Indonesia and the study region.

Figure 1. Location of the study area in Central Lombok Regency, West Nusa Tenggara Province, Indonesia, showing the sub‑districts where swamp buffalo were sampled

Animals and sampling

Sixty adult swamp buffalo owned by smallholder farmers in Central Lombok were included in the study. Animals were selected using purposive sampling based on age, health status and the willingness of owners to participate. Age was estimated from dentition, focusing on the eruption and replacement pattern of permanent incisors. The classification followed a dentition scale described for domestic buffalo Bubalus bubalis, in which the sequential appearance of the first, second, third and fourth pairs of permanent incisors is used to distinguish young adults from older adults, with approximate age ranges adapted to this species (Santos et al 2008). Although the general pattern of incisor eruption is broadly similar to that of river buffalo, swamp buffalo and cattle, the age classes applied in the present study were therefore based on a scale developed for domestic buffalo rather than specifically for swamp buffalo populations. Based on this scale, animals were grouped into two age classes: 36–48 months (younger adults) and 48–60 months (older adults). Within each age group, animals were further classified by sex (males and females). Only clinically healthy animals without evident deformities were included, and all observations and measurements were made on farm under normal management conditions to reflect typical smallholder practices.

Qualitative traits

Qualitative traits were recorded through direct visual observation in the field under natural daylight conditions. The traits observed were head shape (oval, broad), head hair colour (black, grey, brown, white), horn shape (curved backward, curved upward, straight sideways), neck type (with dewlap, without dewlap), neck collar line (single, double), body hair colour (grey, brown, black), skin colour (black, grey, reddish), leg colour (white, grey, black) and tail hair colour (black-brown, grey, brown). For each trait, the number of animals in each category was counted and expressed as a percentage of the total sample.

Quantitative traits

Quantitative traits recorded were shoulder height, body length, chest circumference, hip height, horn length, tail length and bodyweight. Linear body measurements were taken using a flexible measuring tape with animals standing on a flat, level surface. Shoulder height and hip height were measured from the highest point of the withers or hip to the ground; body length was measured from the point of the shoulder to the pin bone; and chest circumference (heart girth) was measured around the thorax just behind the shoulder.

Live bodyweight was not measured using a scale because weighing facilities are not available in the study area under typical smallholder conditions. Instead, bodyweight was estimated using a field formula based on chest circumference (heart girth) and body length, as commonly applied by local extension officers and farmers for swamp buffalo. For each animal, heart girth and body length were measured and the estimated bodyweight was calculated using the standard tape‑based equation recommended for buffalo in the region. This approach reflects routine on‑farm practice and allows smallholders to estimate live weight when scales are absent. The subsequent analyses therefore evaluated the relationship between these field bodyweight estimates and chest circumference to confirm the usefulness of simple tape measurements as a proxy for live weight under low‑input conditions. The formula used was

where CC is chest circumference in cm, BL is body length in cm, and 10840 is a constant used to normalize the units and calculate body weight in kilograms (Setiawan 2022).

Statistical analysis

Qualitative data were summarised as frequencies and percentages. Quantitative data were analysed using descriptive statistics to obtain the mean and standard deviation for each trait by sex and age group. The coefficient of variation for each quantitative trait was calculated as the standard deviation divided by the mean, expressed as a percentage, to indicate relative variability. Traits with coefficients of variation below about 15 percent were interpreted as relatively homogeneous, while values above this level indicated greater variability in the population. Analyses were performed using spreadsheet and statistical software.

Results and discussion

Qualitative phenotypic traits

Adult swamp buffalo in Central Lombok showed a clear dominance of several qualitative traits (Table 1). Most animals had an oval head shape and there were relatively few with broad heads. Head hair was mainly black and brown, with grey animals less frequent and white rare. Backward-curved horns were the most common horn type, followed by upward-curved and straight-sideways horns. All animals observed lacked a dewlap. The neck collar line was mostly double rather than single. Body hair colour was dominated by grey, followed by brown and black, and skin colour was mainly black. Leg colour was predominantly white and tail hair was most often black-brown.

Adult swamp buffalo in Central Lombok typically showed oval head shape, backward-curved horns, grey body hair, black skin and white legs (Figure 2). This combination of traits is consistent with descriptive reports of swamp buffalo in other parts of Indonesia and with general descriptions of indigenous buffalo in tropical environments which often highlight dark or grey coat colours and robust conformation (Prihandini et al 2023; Putri et al 2022; Suhardi et al 2022). The dominance of a few phenotypic categories and the absence of dewlap suggest a relatively uniform external appearance within the population. Recent genomic studies have shown that coat colour variation in swamp buffalo, including the white phenotype, can be explained by specific LINE-1 insertions in the ASIP gene, illustrating the genetic basis of qualitative traits (Nguyen et al 2025; Rehman et al 2021; Trigo et al 2021).

Figure 2. Typical phenotype of adult swamp buffalo in Central Lombok showing oval head shape, backward‑curved horns, grey body hair, black skin and white legs

The dominance of a few qualitative categories and the clear typical appearance can facilitate visual identification of Central Lombok swamp buffalo by farmers and extension workers and may help maintain locally adapted types in smallholder systems. For smallholders, such easily recognisable traits are important for avoiding uncontrolled crossbreeding that might dilute desirable adaptive characteristics such as tolerance to local feed resources, climate and working conditions (Vanvanhossou et al 2025; Widyas et al 2022).

Quantitative phenotypic traits

Quantitative measurements showed that older adult buffalo in the 48–60 month age group had larger body dimensions than younger adults in the 36–48 month group for both sexes. In the older group, mean shoulder height, body length, chest circumference and hip height were all higher and mean bodyweight was greater in males than in females, reflecting sexual dimorphism and continued growth up to about five years of age (Table 2). Bodyweight increased with chest circumference in both males and females, confirming that chest girth can be used as a simple field indicator of bodyweight as reported in other buffalo populations (Aritonang et al 2024; Chen et al 2022; Nastiti et al 2023). Mean shoulder height, body length and chest circumference were all higher in males than in females, which agrees with findings for other indigenous buffalo and reflects the typical sexual dimorphism in body size (Alvin et al 2025; Aritonang et al 2024).

Coefficients of variation for most quantitative traits were below 15 percent, which indicates relatively homogeneous body size within the population. In contrast, horn length and tail length, especially in females, showed higher variability. The body size observed in adult swamp buffalo in Central Lombok falls within the medium range reported for indigenous swamp buffalo in Indonesia and neighbouring countries where mature bodyweights commonly lie between about 350 and 500 kg depending on age and management (Fajar et al 2023; Nastiti et al 2023). Comparative data from other Indonesian buffalo populations such as those from Banten and West Lombok indicate similar patterns of morphometric diversity with body length and chest circumference being key discriminating traits (Nafiu et al 2025; Rusdin et al 2022; Saputra and Anggraeni 2023).

The positive association between bodyweight and chest circumference in this study confirms that simple tape measurements can be used as a practical proxy for live weight estimation in the field, especially where weighing scales are not available (Şengül and Çelik 2025; Soliman 2021). This is particularly useful for smallholder farmers who need to decide on sale, feeding and work allocation based on animal size under low-input conditions. In this context, the relatively homogeneous body dimensions indicate that chest circumference and body length can support straightforward on-farm selection for desired body size and condition without sophisticated equipment.

The combination of uniformity in major body measurements and higher variability in horn and tail length suggests that selection could focus on production-related traits such as body size and weight without greatly altering the typical conformation that farmers value for draught and cultural functions. The phenotypic baseline documented here can serve as a reference for future breeding or conservation programmes aiming to maintain the characteristic phenotype of Central Lombok swamp buffalo while improving productivity under smallholder conditions (Brocklebank et al 2025; Putri et al 2022; Suhardi et al 2022). It can also support efforts to recognise and promote these animals as a distinct local genetic resource within national livestock development strategies.

These results underline the role of locally adapted swamp buffalo as multifunctional animals in integrated smallholder farming systems. Their moderate but robust body size, adaptation to local feed resources and role in providing draught power and manure support sustainable use of local resources and recycling of biomass in mixed crop–livestock systems (Chiariotti et al 2025; Pineda et al 2021). Recent reviews have emphasised the importance of conserving indigenous buffalo genetic resources in Southeast Asia as these animals are well adapted to hot, humid climates and low-input management and can contribute to food security and climate resilience under changing environmental conditions (Ape et al 2025; Pineda et al 2021).

The relatively homogeneous body size and the dominance of a specific qualitative phenotype in Central Lombok indicate that the swamp buffalo kept by smallholder farmers constitute a locally adapted genetic group. At the same time, previous work has shown that swamp buffalo, despite having a consistent overall phenotype, can exhibit somatic and physiological differences depending on climate and land conditions, largely as a result of herd isolation (Minervino et al 2020). This suggests that the Central Lombok population may carry local adaptations that warrant targeted conservation measures before extensive crossbreeding or uncontrolled movements of animals erode its distinctive characteristics.

The phenotypic baseline generated in this study can be used by local authorities, extension services and farmer groups to develop practical guidelines for the conservation and propagation of these buffalo. For example, the dominant phenotype characterised by oval head shape, backward‑curved horns, grey body hair, black skin and white legs can serve as a visual standard when selecting breeding males and females on farm. Simple linear body measurements, particularly chest circumference, body length and shoulder height, can be routinely recorded by farmers and extension workers to identify breeding stock with desirable body size and conformation within the ranges observed in this study. Community‑based breeding programmes that prioritise males with typical phenotypes and adequate body size and promote their use across farmer groups could help maintain local adaptation while gradually improving productivity under smallholder conditions.

In addition, the phenotypic information provided here can serve as a starting point for more detailed genetic characterisation of Central Lombok swamp buffalo. Future research could combine phenotypic monitoring with molecular tools such as mitochondrial markers and genome‑wide SNP data to identify unique genetic signatures of this population, quantify genetic diversity and clarify relationships with other Indonesian swamp buffalo. Such genotypic evaluations would allow the development of region‑specific breeding and conservation strategies, complementing the phenotypic descriptors documented here and supporting the recognition of Central Lombok swamp buffalo as a distinct local genetic resource within national livestock development policies.

Conclusions

Adult swamp buffalo kept by smallholder farmers in Central Lombok, Indonesia, show a dominant phenotypic profile characterised by oval head shape, backward‑curved horns, grey body hair, black skin, white legs and black‑brown tail hair. Quantitative measurements indicate that older adult animals have larger body dimensions and higher bodyweight than younger adults and that males are larger than females, with most body traits showing relatively low variability while horn and tail length are more variable. These features suggest that the local buffalo population has a relatively uniform body size and a characteristic external appearance, which is advantageous for on‑farm identification and selection of breeding animals in smallholder systems. The strong association between chest circumference and field‑estimated bodyweight confirms that simple tape measurements can be used by farmers to estimate live weight where scales are not available, supporting practical decisions on sale, feeding and work allocation. The phenotypic baseline provided by this study offers a useful reference for the management and conservation of local swamp buffalo genetic resources and can inform community‑based selection schemes that use visual and linear body traits as criteria for choosing breeding males and females. In the longer term, integrating these phenotypic descriptors with genetic evaluations will be essential to design region‑specific breeding and conservation programmes that maintain the characteristic phenotype and adaptive traits of Central Lombok swamp buffalo while improving their productivity in mixed crop–livestock farming systems.

Acknowledgments

The authors gratefully acknowledge the smallholder farmers in Central Lombok who allowed access to their animals and shared information on their management practices. We thank the staff and students of the Laboratory of Animal Breeding and Genetics, Faculty of Animal Science, University of Mataram, for their assistance with field data collection and animal handling. This study was supported by internal research funding from the University of Mataram and the Faculty of Animal Science. The authors also appreciate the constructive comments from colleagues that helped improve the design and interpretation of this study.

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