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Article

Ecological and Population Dynamics of Black and Grey Francolin (Phasianidae) in Totali Game Reserve, Pakistan

by
Asad Ullah
1,*,
Sumaira Shams
1,*,
Sultan Ayaz
2,
Eliana Ibáñez Arancibia
3,4,5,
Unays Siraj
6,
Patricio R. De los Rios-Escalante
5,7,8,
Taqi Ullah
2 and
Farhad Badshah
1,9,10
1
Department of Zoology, Abdul Wali Khan University, Mardan 23200, Pakistan
2
College of Veterinary Sciences and Animal Husbandry, Abdul Wali Khan University, Mardan 23200, Pakistan
3
PhD Program in Sciences Mentioning Applied Molecular and Cell Biology, La Frontera University, Temuco 4780000, Chile
4
Laboratory of Engineering, Biotechnology and Applied Biochemistry—LIBBA, Department of Chemical Engineering, Faculty of Engineering and Science, La Frontera University, Temuco 4780000, Chile
5
Departamento de Ciencias Biologicas y Químicas, Facultad de Recursos Naturales, Universidad Católica de Temuco, Casilla 15-D, Temuco 4780000, Chile
6
Department of Biotechnology, Faculty of Science, Mahidol University, Bangkok 10400, Thailand
7
Núcleo de Estudios Ambientales UC Temuco, Casilla 15-D, Temuco 4780000, Chile
8
Instituto de Estadística, Facultad de Ciencias, Pontificia Universidad Católica de Valparaíso, Av. Brasil 2830, Valparaíso 2340025, Chile
9
Shenzhen Branch, Guangdong Laboratory for Lingnan Modern Agriculture, Key Laboratory of Livestock and Poultry Multi-Omics of MARA, Agricultural Genomics Institute at Shenzhen, Chinese Academy of Agricultural Sciences, Shenzhen 518124, China
10
State Key Laboratory of Animal Biotech Breeding, Institute of Animal Sciences, Chinese Academy of Agricultural Sciences, Beijing 100193, China
*
Authors to whom correspondence should be addressed.
Birds 2025, 6(2), 26; https://doi.org/10.3390/birds6020026
Submission received: 4 April 2025 / Revised: 13 May 2025 / Accepted: 15 May 2025 / Published: 22 May 2025
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Simple Summary

This study examines the ecological factors influencing black and grey francolin populations in the Totali Game Reserve, Pakistan. We found significant density variations using distance sampling and vegetation assessments across three habitat types—agricultural lands, agroforests, and pasturelands—with both species favoring pastureland ecosystems. Population peaks occurred in autumn due to post-breeding surges and optimal habitat conditions. The birds strongly preferred areas dominated by key plant species such as hopbush, raspberry, barberry, and blackberry, which provide essential resources for foraging, nesting, and shelter. This study underscores the need for conservation efforts to preserve pasturelands and to implement adaptive management strategies to counter population declines.

Abstract

Ecological determinants of avian population dynamics are essential for effective wildlife management and conservation. Black and grey francolins are classified as being of least concern by the IUCN and are an ecologically significant bird in the northern regions of Pakistan. Although habitat degradation and overhunting have adversely affected their populations, research on the population structure and habitat preferences of the Totali Game Reserve, Buner District, Pakistan, remains limited. The present study applied distance sampling via the line transect method to estimate population densities and the quadrat method to assess vegetation characteristics across three distinct habitats. A statistical analysis revealed significant variations in densities across the three habitat types—agricultural lands, agroforests, and pasturelands—with both species exhibiting distinct preferences for pastureland ecosystems. Seasonal trends indicated that autumn represented the peak period for the population, driven by post-breeding surges and optimal habitat conditions. Moreover, both species demonstrated a strong affinity for habitats dominated by specific plant species, including hopbush, raspberry, barberry, and blackberry, which provide a critical resource for food, foraging, nesting, and shelter. These findings highlight the ecological importance of pasturelands and specific vegetation types for supporting the survival of these species. Furthermore, conservation efforts should prioritize the preservation and restoration of pastureland ecosystems and incorporate rigorous monitoring programs to guide adaptive management strategies aimed at mitigating population declines.

1. Introduction

Regional biodiversity continues to decline as human activities intensify across natural communities, leading to critical environmental issues [1]. Birds are a crucial indicator of ecosystem health [2]. Their presence or absence reflects biodiversity across ecological communities, offering insights into environmental conditions and the vitality of various habitats [3]. Species richness and the diversity of landscapes not only provide valuable information in avian spatial ecology but also highlight the significance of habitat structure in sustaining ecological processes [4,5]. In recent years, research exploring the connection between birds and vegetation has taken place primarily in natural landscapes [1,6,7]. However, with rapid urbanization, studying these relationships within protected areas has become increasingly vital [8,9].
Among avian groups, Galliformes represent one of the most ecologically and culturally significant orders. Galliformes species are primarily maintained through long-term association with human societies through domestication, subsistence hunting, and their incorporation into traditional trade [10,11,12]. The order Galliformes comprises 295 species across five families, including Phasianidae, which features chickens, pheasants, and francolins [12,13]. Globally, 41 species of francolins have been identified, including five native to the Indian subcontinent, including the black (Francolinus francolinus) and grey francolin (Francolinus pondicerianus) [14]. Birds of the francolins’ genus (Francolinus) are medium-sized birds that rely on the ground for shelter and sustenance [15,16]. According to the International Union for Conservation of Nature’s (IUCN) Red List, both species are currently classified as least concern species due to their wide geographic distributions and large global populations [17]. Further, black and grey francolins exhibit a partially overlapping global distribution. Black francolins predominantly inhabit moist lowland habitats, ranging from Cyprus to Northeast India, whereas grey francolins are typically associated with the semi-arid plains extending from Sri Lanka to the Persian Gulf [18,19]. In Pakistan, they are predominantly found in undisturbed tropical thorn forests, the Indus plan, and the lower hills of Makran and Lasbella (Balochistan), as well as protected areas of Cherat [20,21,22]. Additionally, these species are observed in protected areas like the lower hills of Kohat, Salt Range, Potohar plateau [20,23], and Buner regions [24]. The grey francolin predominantly inhabits open cultivated pasturelands but is rarely recorded above 1200 m elevation [25]. In contrast, the black francolin occupies a broader elevation range (300–2100 m), is widely distributed across Asian states, and held in particularly high esteem by local farmers [26,27]. Both francolin species play crucial roles in agriculture and forest ecosystems by consuming a variety of insects [20]. They forage in pasturelands, agricultural lands, and forests, consuming seeds, insects, shoots, ants, and caterpillars [22].
The habitat-specific distribution patterns of these birds are influenced by food availability, vegetation structure, and seasonal changes [28]. Pasturelands and agroforests are particularly vital, as they provide essential resources for nesting, foraging, and shelter [29]. Previous studies in the Salt Range and Pothwar Plateau in Pakistan have demonstrated that bird populations are strongly linked to vegetation composition and habitat characteristics [26,30]. A recent survey in Mang Game Reserve, Haripur, reported a density of 0.6 to 4.4 individuals / km 2 [29], with protected areas such as Lehri Park and Lal Suhanra National Parks supporting higher densities of approximately 6 to 8 individuals / km 2 , respectively [20,31].
Vegetation compositions and food availability are key determinants of francolin population dynamics, as these birds depend on natural habitats for foraging and nesting [32,33]. Positive associations between plant species diversity and francolin density emphasize the importance of vegetation structure in determining habitat suitability [34]. Seasonal variation further highlights the need for temporal habitat management, as species densities peak in post-breeding autumn when environmental conditions and food availability are optimal [27]. However, anthropogenic stressors, such as the misuse of agrochemicals, industrial pollutants, and alterations in temperatures and precipitation patterns, have significantly disrupted habitat quality, spatial distribution, and reproductive success in these species [35,36]. These environmental pressures contribute to impaired growth rates, reduced body mass, and increased susceptibility to diseases [37,38].
Historically, recognized as game birds, francolins have held deep cultural significance in Pakistan, often representing social status and traditional values within local communities. Francolins have faced increasing pressure from hunting [22], deforestation [23,39], agricultural expansion [40], and urbanization [7], all of which contributed to a sharp population decline in recent decades. However, black and grey francolins are legally protected in designated regions of Pakistan, with controlled hunting permitted during the non-breeding season [41]. Despite their ecological importance, research on the population dynamics and habitat preferences of black and grey francolin remains limited. These research gaps are particularly evident in protected areas such as the Totali Game Reserve, a key conservation site in northern Pakistan. This study aims to address four key questions: (1) Do black and grey francolin populations differ across habitat types within the study area? (2) Do their populations vary seasonally across these habitats? (3) Which habitats are mostly preferred by these species? (4) Do plant species occurring in various habitats positively or negatively affect their populations? We hypothesize that the populations of black and grey francolin in the Totali Game Reserve vary significantly across habitat types and seasons. We further predict that specific habitats and their associated plant species have measurable positive or negative effects on the abundance and habitat preferences of these species. This study provides a foundation for establishing conservation priorities and sets the stage for future long-term studies on francolin ecology.

2. Materials and Methods

2.1. Study Area

We conducted our research in the Totali Game Reserve, Pakistan, a protected wildlife zone covering 30,960 ha, with altitudes ranging from 500 to 2100 m above sea level (Figure 1). This study focused on black and grey francolin and was carried out from October 2022 to September 2023. The reserve, located at 34° 25 ′ 29.14 ″ N, 72° 39 ′ 56.83 ″ E, features diverse habitat types characterized by varied vegetation and climatic conditions [24,42]. The region experiences a seasonal climate: winter extends from November to March, the dry season spans June to September, and monsoon rains occur from July to August. The average annual rainfall in the study area was 76.5 mm. The highest average temperature (31.5 ± 1.7 °C) was recorded during the dry season, while the lowest occurred in winter (1.7 ± 2.9 °C) (Table S1) [43]. Based on a pre-survey conducted from June to August 2022, the study area was classified into three habitat types (Figure 1). During this pre-survey, systematic walks were carried out across multiple sites to collect baseline data on vegetation composition, land use, and the presence of key species: 1. agricultural lands, which include cultivated crops, pastures, and lands interspersed with natural vegetation; 2. pasturelands, dominated by herbaceous plants and shrubs; and 3. agroforests, characterized by rugged terrain with significant elevation changes featuring tall trees, dense ground cover, and upper vegetation layers with open patches dominated by grasses. The surveyed data were assumed to represent the reserve-wide patterns to better understand the species’ habitat and population dynamics [44].

2.2. Identification of Preferred Habitats

The quadrat method was employed to identify the preferred habitats of the two francolin species during October 2022 to September 2023, across three habitat types, agricultural lands, agroforests, and pasturelands, by analyzing the vegetation composition within the study area. Quadrat sampling is a widely used technique in ecological research, particularly for assessing plant biodiversity and community structure [20,45]. It involves the direct counting of organisms within a defined sample area, providing quantitative data for vegetation analysis [45,46]. Briefly, three quadrats were randomly placed in the middle of each transect within the three habitat types to ensure representative sampling. The quadrat size was standardized based on vegetation type: trees were sampled in 10 m × 10 m, shrubs in 5 m × 5 m, and herbs in 1 m × 1 m quadrats following established protocols [29].

2.3. Density Assessment

To determine the densities of black and grey francolin, the distance sampling method, specifically the line transect method, was utilized. Briefly, to obtain density estimates by adjusting the data to account for species detectability [22], this method considers the varying probability of detecting bird species based on the distance from the observer [18,26]. Twelve transects, each measuring 1000 m in length and 50 m in width, were randomly established across the three vegetation habitats, serving as the primary unit for bird surveys (Figure 1). The survey area extended 25 m on either side of the transect line, within which individuals of each species were recorded [27,47]. To ensure spatial independence, a minimum separation of 100 m was maintained between adjacent transects, and their distribution was arranged as evenly as possible [27]. However, some transects were separated by distances of several kilometers. Surveys during these trials were conducted twice daily: at dawn (6:00–8:00 am) and dusk (4:00–6:00 pm). The means of both sessions were used to estimate the population densities for each transect. The total survey time per trial, combining both dawn and dusk sessions, ranged from two to three hours. Because of the significant distance between the survey sites, the first round of sampling of each site was conducted during the first two weeks of the month, followed by the second round in the latter half. To maintain temporal consistency, survey sites were visited in the same sequence each month, ensuring data collection within comparable time frames. Data collection focused on detected individuals, recording their numbers, detection cues, observer position, and distance to the species. Birds flying overhead were excluded from the counts. Species densities were calculated per hectare and averaged monthly to analyze seasonal trends using detection probabilities based on the observer distance. Two trained observers conducted the surveys while walking at 1–2 km/h to ensure methodological consistency. The densities of the two species were calculated using the following formula [22,41]:
D = Σ n 2 L W
where L = Total length of the transect, n = the number of birds detected, W = Width of the transect, and D = Estimated density of the species.

2.4. Statistical Analysis

The means and standard deviations (mean ± SD), and transect data were calculated to summarize the density variations. Before the analysis, we assessed the normality of the data using the Shapiro–Wilk test [48]. As the data were normally distributed, one-way ANOVA was applied to determine the statistical significance of the differences in the densities among habitats and seasons. Paired sample t-tests were used to evaluate the interspecific differences within each habitat. To further analyze the relative density variation in more detail, we performed a similarity percentage analysis (SIMPER), a cluster analysis, and an analysis of similarities (ANOSIM) using the Bray–Curtis quantitative similarity on species detections from sampled transects. To show the relationship between the species and preferred plant species, a generalized linear model (GLM) with a Gaussian distribution was utilized. This approach enabled the evaluation of a potential association between the two bird species and plants across the study area, providing deeper insights into the ecological interactions that shape these relationships. All statistical analyses were conducted using R software (V. 4.34).

3. Results

3.1. Overview

Over the study period, a total of 3014 individuals from both species were observed. Grey francolin was the dominant species, at 60.71% (n = 1830), while black francolin accounted for 39.29% (n = 1184). The black francolin numbers were highest in the agroforests (101.8 ± 3.5, n = 509), followed by agricultural lands (98.33 ± 5.85, n = 295) and pasturelands (95 ± 8.98, n = 380). However, no statistically significant variation in black francolin abundance was found among the three habitats (F = 1.29, df = 2, p > 0.05). In contrast, grey francolin was most abundant in the pasturelands (171 ± 4.24, n = 684), followed by agroforests (127.6 ± 13.24, n = 638) and agricultural lands (169.33 ± 12.89, n = 508). A statistically significant difference in grey francolin distribution across these habitats was detected (F = 22.002, df = 2, p < 0.001).

3.2. Population Density of Black and Grey Francolin

The densities of black and grey francolin across three distinct habitats were assessed using data from the twelve transects. For black francolin, the highest density was recorded in transect 6 (2.7, 1.77 ± 0.59), followed closely by transect 5 (2.7, 1.76 ± 0.56) (Figure 2a). The lowest density was recorded in transect 4 (2.30, 1.29 ± 0.59), located in pasturelands (Figure 2a). Similarly, transect 6 also showed the highest density of grey francolin (4.18, 3.02 ± 0.8), followed by transect 7 (4.37, 2.93 ± 0.79), both located in pasturelands. In contrast, the lowest density was recorded in transect 11 (2.91, 1.66 ± 0.60), within the agroforests (Figure 2b).
Paired t-tests revealed significant differences in density between black and grey francolins across all three habitat types. In agricultural lands, black francolin had a mean density of 1.58 individuals/ha, while grey francolin exhibited a significantly higher density of 2.73 individuals/ha (t = 9.12, df = 11, p < 0.001) (Figure 3). In pasturelands, the mean density of black francolin was 1.60 individuals/ha, while grey francolin exhibited a slightly higher density of 1.82 individuals/ha (t = 10.14, df = 11, p < 0.001) (Figure 3). In the agroforests, black francolin had a significantly lower mean density of 1.54 individuals/ha, whereas grey francolin exhibited a significantly higher density of 2.01 individuals/ha (t = 4.45, df = 11, p < 0.001) (Figure 3).
Further, an analysis of similarity (ANOSIM) revealed significant dissimilarity among transects across all habitat types (global R = 0.629, p < 0.001). In contrast, a SIMPER analysis showed varying levels of similarity between transects from pasturelands (40.88 %), agricultural lands (32.14 %), and agroforests (26.98 %) (Figure 4). These findings revealed notable spatial variations in the densities of grey and black francolin within the Totali Game Reserve.

3.3. Seasonal Variation in Population Density

We further investigated seasonal variation in the densities of black and grey francolin. The results revealed significant variation in densities among the various seasons, with both species exhibiting the highest density during autumn (Figure 5). Black francolin exhibited significantly higher densities in autumn (F = 16.63, df = 3, p < 0.001), with an average density (2.85, 1.90 ± 0.63), while the lowest density occurred in the summer (2.22, 1.22 ± 0.38). Similarly, grey francolin reached its peak density in the autumn (F = 18.36, df = 3, p < 0.001), with a mean density (4.37, 3.12 ± 0.65), whereas the lowest densities were recorded in summer (3.29, 2.08 ± 0.60). Average densities were observed for both species during spring and winter.

3.4. Habitat-Specific Associations

A generalized linear model was applied to assess the relationship between targeted bird species and the surrounding plant community. Hopbush (Dodonea viscosa), barberry (Berberis lycium), and raspberry (Rubus ellipticus) showed positive associations with bird densities, indicating that these plants are likely to provide important resources. A strong positive association of black francolin was observed with hopbush (t = 27.14, df = 11, p < 0.05), raspberry (t = 26.89, df = 11, p < 0.05), and barberry (t = 16.95, df = 11, p < 0.05) (Table 1). Although fleabane (Conyza Canadensis), marijuana (Cannabis sativa), kamala tree (Mallotus philippensis), and blackberry (Rubus fruticosus) showed positive trends, their relationships were not statistically significant. Similarly, negative but non-significant associations were observed with toothache tree (Zanthoxylum aromatum), chir Pine (Pinus roxburghii), and phulai (Acacia modesta) plant species. In contrast to black francolins, the density of grey francolins did not show a significant association with the observed plant species. Although some plant species such as raspberry (t = 1.81, df = 11, p > 0.05), hopbush (t = 1.03, df = 11, p > 0.05), and barberry (t = 0.51, df = 11, p > 0.05) showed a slight positive association with grey francolin density (Table 2). Similarly, non-significant negative trends were observed for fleabane, marijuana, toothache tree, and chir pine. The GLMs suggested that certain plant species might enhance habitat suitability, while others might reduce it.

4. Discussion

Our study revealed distinct density estimates for black and grey francolin in the Totali Game Reserve based on a year-long population assessment. Overall, the average highest density for both francolin species was found in the pasturelands. Generally, our recorded values appear to be lower compared to prior reported studies on black and grey francolins in the Mang Game Reserve and Lal Suhanra National Park, Pakistan [29,41]. Research that took place in various parts of Pakistan and India reported population densities ranging from 1 to 8 individuals/ha [20,25,27,29]. We highlight that the focus on laying transects in the various regions of the study area is likely to provide an accurate density value [49]. Prior studies on black francolin in the Kohat region and Lal Suhanra Park of Pakistan have reported significantly higher densities [22,50]. In comparison to our findings, higher densities have been reported both in wetlands [29,31] and in agroforests [51].
Similarly, grey francolin exhibited higher densities both in agroforests [29] and in pasturelands [25], as well as in Lal Shanara Park [51]. These densities were generally higher than those observed in our study, suggesting possible regional or habitat-specific variations in population dynamics. Differences in environmental conditions, resource availability, and human disturbance likely contribute to such discrepancies [52,53]. As for habitat, this study reveals significant partitioning of black and grey francolin, with differences in abundance observed across transects spanning agricultural lands, pasturelands, and agroforests. The two species replaced each other within these habitats but showed distinct density patterns throughout the surveyed area. These quantitative results provide strong empirical support for these preferences, reinforcing trends reported in previous research [22]. However, the observed replacement dynamics and density variations across habitats highlight the need for further investigation at larger scales. Understanding the ecological drivers behind these patterns, such as competition [54], resource availability, and environmental pressure [55,56], can play an essential role in refining conservation strategies [31,35].
In game-related bird species, habitat significantly influences species distribution [57], abundance, behavior [58], and lifespan [30]. Additionally, wildlife distribution is often uneven and fluctuates across time and seasons [59]. Habitat selection is primarily driven by the availability of food and shelter [60]. Studying and understanding population dynamics and habitat preferences is essential for developing effective conservation strategies [20]. This study revealed that black and grey francolin exhibited distinct habitat preferences, with higher densities in pasturelands dominated by key plant species such as hopbush and raspberry. Notably, the autumn season emerged as the peak season for both species’ populations, driven by post-breeding increases and optimal environmental conditions. Consistently elevated densities throughout all seasons imply that these regions provide optimal habitat conditions. The post-breeding period, during which juvenile individuals fledged, contributes to this population rise [24]. Moreover, abundant food resources, such as seeds and fruits, attract these species, while the mild autumn temperatures further facilitate their abundance.
The current study highlighted notable habitat-specific differences in the population densities of black and grey francolin, reinforcing previously observed ecological trends [29]. The cluster analysis separated agroforest transects (T8, T9, T10, and T12) from those in agricultural lands (T6 and T7) and pasturelands (T2 and T1), indicating consistent species detection within each habitat type. However, altered transects T3 (agricultural lands) and T5 (pasturelands) suggested the presence of a transitional zone between habitat types. These findings, supported by ANOSIM and SIMPER results, revealed significant dissimilarities in species composition between the three habitats. Furthermore, grey francolin showed significantly higher population densities along transects T1, T4, and T7, situated in agricultural lands and pasturelands. These observations align with previous reports showing higher grey francolin densities in plain areas [61], pasturelands [29], and wetland-adjacent wild areas [34]. Similarly, black francolin exhibited higher densities along pastureland transects (T6 and T7). However, the literature supports that black francolin prefers more humid environments such as wetlands [20], moist pasturelands [27], and mountainous forests associated with ecotones [51]. These habitat preferences explained the high densities of grey francolin in more open and drier habitats, where black francolin appears to be less dominant. The adaptability of grey francolin to open pasturelands and cultivated fields, coupled with its generalist diets of seed, insects, and vegetation [61], allows it to exploit a broader range of habitats and food resources [3,33].
Ecological adaptability enables broader geographic distribution and supports higher population densities in habitats characterized by low vegetation cover [51]. Habitat and ecological differences, such as food availability, can shape the reproductive strategies of black francolin [24]. However, the grey francolin was more frequently recorded for short distances, likely due to its preference for habitats with dense vegetation cover. Importantly, our study revealed the vegetation’s role in supporting the two species’ populations and their complex interdependencies between the associated habitat and avian ecology. A diverse range of plant species fulfilled their essential food and protection. Grey francolin tends to have branched roots aligned with its known habitat associations [30]. Ground vegetation in the pasturelands contributed to the high densities observed for both species, while lower densities were observed in the agroforests. These findings indicate that pasturelands provide optimal conditions for these species. The results are consistent with those in the existing literature indicating that black and grey francolin prefer specific shrub species, such as hopbush, raspberry, barberry, and blackberry for perching and roosting, showing a direct relationship with vegetation structure that influences habitat selection [20,29]. Our findings align with those of previous studies that vegetation between 4 and 48 cm in height supports greater bird species diversity compared to shorter vegetation [62]. However, the compounded challenges driven by urbanization and hunting [2,4,63] potentially declined francolin species and their synergistic diversity [64,65]. Grey francolin is more vulnerable to altered habitats and has demonstrated a decline in density due to decreased habitat availability [26,34].
This study highlights the need for conservation efforts beyond ecological factors, emphasizing the importance of coordinated strategies to restore and prioritize habitats essential for sustaining francolin populations. These findings also support a broader scientific approach for seasonal habitat selection and vegetation dynamics in avian species [49]. However, our research is limited by its spatial scope, lack of genetic data, and information on reproductive success, predation rates, and gradients of anthropogenic disturbance. Moreover, the present study was conducted over one year and restricted to the Totali Game Reserve, District Buner, Pakistan, which may limit the broader applicability and generalizability of the findings to other regions or habitat types. In addition, the absence of tracking-based approaches restricts insights into fine-scale habitat use and movement patterns. Future research can integrate long-term monitoring across multiple reserves to capture inter-annual variability and broader ecological patterns. Additionally, molecular tools such as microsatellite or genomic analysis are recommended to assess population structure and gene flow, especially in sympatric zones.

5. Conclusions

Our study elucidates the ecological determinants shaping the population dynamics and habitat selection of black and grey francolin within the Totali Game Reserve of Pakistan. Our study marked spatial and temporal variability in both species’ densities, with the pasturelands showing the highest population concentration, particularly, in the post-breeding autumn period. The preferred plant species of both bird species were hopbush, raspberry, barberry, and blackberry, which are vital for foraging and nesting activities across the three habitat types. These findings reinforce the significance of habitat heterogeneity and plant structural complexity in supporting avian biodiversity. Nonetheless, anthropogenic disturbance, including habitat fragmentation, urban encroachment, and unsustainable land use, significantly pose threats to the persistence of these species. Therefore, it is imperative to integrate ecological monitoring with molecular tools such as genome-wide association studies or microsatellite analyses to further assess genetic diversity, population connectivity, and adaptive potential. Based on advanced analyses, the evolutionary history, and the ecological adaptability of these species, the long-term conservation of galliformes can be potentially enhanced in both natural and anthropogenically altered areas.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/birds6020026/s1, Table S1: Long-term climate data on the Totali Game Reserve, Buner district of Khyber Pakhtunkhwa, Pakistan (October 2022–September 2023). Coordinates: (34° 25 ′ 29.14 ″ ; 72° 39 ′ 56.83 ″ ; altitude: 2100 m asl).

Author Contributions

A.U.: Formal analysis, Investigation, Methodology, Visualization, and Writing—original draft. S.S. and E.I.A.: Conceptualization, Supervision, Methodology, Funding acquisition, Project administration, and Writing—review & editing. Conceptualization, Supervision, Methodology, Funding acquisition, Project administration, and Writing—review and editing. S.A. and U.S.: Conceptualization, Data curation, Formal analysis, Investigation, Visualization, and Writing—review and editing. P.R.D.R. and T.U.: Validation, Data curation, and Writing—review and editing, and F.B.: Resources, Funding acquisition, Formal analysis, Validation, and Writing—review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

This study was financially supported by Project MECESUP UCT 0804.

Institutional Review Board Statement

All research procedures were approved by the Institutional Ethics Committee of the Animal Ethics and Welfare Committee of the Department of Zoology, Abdul Wali Khan University, Mardan, Pakistan (PK) (approval No. AWK-ZOO24906), and all methods were carried out according to the Code of Practice for the Care and Handling of Animal Guidelines.

Data Availability Statement

The data from this study are available from the corresponding authors upon reasonable request.

Acknowledgments

Our research team sincerely thanks the District Forest Officer (Buner Wildlife Division) and staff for their valuable support. We especially appreciate the experts of the Totali Game Reserve, District Buner, who provided us with all the relevant information during the site visits.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Map of the Totalai Game Reserve showing the locations of transects (T1–T12), streams, roads, and contour lines. Transects are color-coded by habitat type: agricultural lands (stratum a) in blue, pasturelands (stratum b) in green, and agroforests (stratum c) in black. The coordinates for each transect are listed on the left side of the map.
Figure 1. Map of the Totalai Game Reserve showing the locations of transects (T1–T12), streams, roads, and contour lines. Transects are color-coded by habitat type: agricultural lands (stratum a) in blue, pasturelands (stratum b) in green, and agroforests (stratum c) in black. The coordinates for each transect are listed on the left side of the map.
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Figure 2. Transect-wise population density (individuals/ha; mean ± SD) of (A) black and (B) grey francolin across the three vegetation habitats. Stratum A (in green): agricultural lands; stratum B (in orange): pasturelands; stratum C (in grey): agroforests.
Figure 2. Transect-wise population density (individuals/ha; mean ± SD) of (A) black and (B) grey francolin across the three vegetation habitats. Stratum A (in green): agricultural lands; stratum B (in orange): pasturelands; stratum C (in grey): agroforests.
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Figure 3. Violin plots with overlaid boxplots representing the densities (individuals/ha) of black francolin (blue) and grey francolin (green) across each habitat. The significance of any observed differences (p < 0.05) was assessed using a paired sample t-test.
Figure 3. Violin plots with overlaid boxplots representing the densities (individuals/ha) of black francolin (blue) and grey francolin (green) across each habitat. The significance of any observed differences (p < 0.05) was assessed using a paired sample t-test.
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Figure 4. Cluster analysis using the Bray–Curtis quantitative similarity measure on the detection of black and grey francolin in the different transects sampled. Dendrogram including all transects (T1–T12). Stratum A (in yellow): agricultural lands; stratum B (in blue): pasturelands; stratum C (in green): agroforests.
Figure 4. Cluster analysis using the Bray–Curtis quantitative similarity measure on the detection of black and grey francolin in the different transects sampled. Dendrogram including all transects (T1–T12). Stratum A (in yellow): agricultural lands; stratum B (in blue): pasturelands; stratum C (in green): agroforests.
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Figure 5. Violin plots with overlaid boxplots representing the seasonal variations in densities (individuals/ha) of (A) black francolin and (B) grey francolin.
Figure 5. Violin plots with overlaid boxplots representing the seasonal variations in densities (individuals/ha) of (A) black francolin and (B) grey francolin.
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Table 1. Parameters of the generalized linear models (GLMs, with Gaussian distribution) testing the relationships between black francolin population density and plant species in the three habitats at the Totali Game Reserve.
Table 1. Parameters of the generalized linear models (GLMs, with Gaussian distribution) testing the relationships between black francolin population density and plant species in the three habitats at the Totali Game Reserve.
Scientific NameCommon NameEstimateStd. ErrorDft ValuePr (>|t|)
Intercept 1.2700.00811161.6900.004
Conyza canadensisFleabane0.0070.001116.6750.095
Dodonea viscosaHopbush0.0420.0021127.1420.024
Cannabis sativaMarijuana0.0060.001116.3860.099
Mallotus philippensisKamala Tree0.0140.003114.7880.131
Berberis lyciumBarberry0.0460.0031116.9500.038
Rubus ellipticusRaspberry0.0390.0011126.8920.024
Rubus fruticosusBlackberry0.0110.0011110.1650.062
Zanthoxylum aromatumToothache Tree−0.0330.00111−28.3000.023
Pinus roxburghiiChir Pine−0.0220.00111−18.9750.034
Acacia modestaPhulai−0.0050.00111−6.6440.096
Table 2. Parameters of the generalized linear models (GLMs, with Gaussian distribution) testing the relationships between grey francolin population density and plant species in the three habitats at the Totali Game Reserve.
Table 2. Parameters of the generalized linear models (GLMs, with Gaussian distribution) testing the relationships between grey francolin population density and plant species in the three habitats at the Totali Game Reserve.
Scientific NameCommon NameEstimateStd. ErrorDft ValuePr (>|t|)
Intercept 3.7610.903114.1640.150
Conyza canadensisFleabane−0.1800.11411−1.5730.361
Dodonea viscosaHopbush0.1820.177111.0300.491
Cannabis sativaMarijuana−0.1010.10111−1.0000.500
Mallotus philippensisKamala Tree−0.4850.32711−1.4800.378
Berberis lyciumBarberry0.1590.310110.5120.699
Rubus ellipticusRaspberry0.3040.167111.8190.320
Rubus fruticosusBlackberry0.0210.128110.1630.897
Zanthoxylum aromatumToothache Tree−0.0510.13411−0.3760.771
Pinus roxburghiiChir Pine−0.2480.13611−1.8190.320
Acacia modestaPhulai−0.1270.09411−1.3600.404
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Ullah, A.; Shams, S.; Ayaz, S.; Ibáñez Arancibia, E.; Siraj, U.; De los Rios-Escalante, P.R.; Ullah, T.; Badshah, F. Ecological and Population Dynamics of Black and Grey Francolin (Phasianidae) in Totali Game Reserve, Pakistan. Birds 2025, 6, 26. https://doi.org/10.3390/birds6020026

AMA Style

Ullah A, Shams S, Ayaz S, Ibáñez Arancibia E, Siraj U, De los Rios-Escalante PR, Ullah T, Badshah F. Ecological and Population Dynamics of Black and Grey Francolin (Phasianidae) in Totali Game Reserve, Pakistan. Birds. 2025; 6(2):26. https://doi.org/10.3390/birds6020026

Chicago/Turabian Style

Ullah, Asad, Sumaira Shams, Sultan Ayaz, Eliana Ibáñez Arancibia, Unays Siraj, Patricio R. De los Rios-Escalante, Taqi Ullah, and Farhad Badshah. 2025. "Ecological and Population Dynamics of Black and Grey Francolin (Phasianidae) in Totali Game Reserve, Pakistan" Birds 6, no. 2: 26. https://doi.org/10.3390/birds6020026

APA Style

Ullah, A., Shams, S., Ayaz, S., Ibáñez Arancibia, E., Siraj, U., De los Rios-Escalante, P. R., Ullah, T., & Badshah, F. (2025). Ecological and Population Dynamics of Black and Grey Francolin (Phasianidae) in Totali Game Reserve, Pakistan. Birds, 6(2), 26. https://doi.org/10.3390/birds6020026

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