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Behavioral Determinants of Mating Success in Blackbuck under Semienclosed Condition: A Cross-Sectional Study in Shuklaphanta National Park, Nepal

Behavioral Determinants of Mating Success in Blackbuck under Semienclosed Condition: A... Hindawi International Journal of Zoology Volume 2022, Article ID 8713116, 7 pages https://doi.org/10.1155/2022/8713116 Research Article Behavioral Determinants of Mating Success in Blackbuck under Semienclosed Condition: A Cross-Sectional Study in Shuklaphanta National Park, Nepal 1 1 2 Puja Saud , Juna Neupane , and Mukesh Kumar Chalise Central Department of Zoology, Institute of Science and Technology, Tribhuvan University, Kathmandu, Nepal Nepal Biodiversity Research Society (NEBORS), Lalitpur, Nepal Correspondence should be addressed to Juna Neupane; zunaneupane@gmail.com Received 1 April 2022; Revised 19 June 2022; Accepted 12 July 2022; Published 5 August 2022 Academic Editor: Irene Pellegrino Copyright © 2022 Puja Saud et al. �is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Knowing reproductive behavior is important for establishing a viable population of nationally endangered blackbuck in Nepal. �is study explored the behavioral events and states associated with copulatory mountings in blackbuck by focal animal sampling method under semienclosed conditions of Hirapurphanta within Shuklaphanta National Park, Nepal. We recorded diˆerent behavioral initiations for the mating reproductive activities and the behaviors followed after copulatory mounting. Four dominant adult males were used as a subject. �e highest number of copulatory mountings was observed during the months of Febru- ary–March. Among the diurnal observational phases, the highest number of copulatory mountings was recorded during 9.00–12.00 hours followed by 15.00 hours onwards. Copulatory mounting was signi“cantly preceded by sni”ng the vulva of female and showing •ehmen response, whereas copulatory mounting was followed by walking. eˆorts in the wild condition, ex-situ breeding in the captivity 1. Introduction and introduction to the semiwild habitats are undergoing. Blackbucks (Antilope cervicapra, Bovidae, Artiodactyla) Captive breeding is one of the best ways to ensure the were formerly widespread across the Indian subcontinent future existence of any plant or animal. However, the captive but are now limited to only small, scattered herds in Nepal, environment can be aˆecting the animal behavior mainly in India, and Pakistan due to excessive hunting and reduced four ways: the regular presence of large numbers of unfa- habitat range [1]. Blackbuck is one of the highly threatened miliar and behaviorally diverse (passive-active) humans, and nationally protected species in Nepal [2]. �ey were restricted and unchanging space, forced proximity to other thought almost extinct until a small herd of nine individuals animal species, and human management [5]. Eˆects of was discovered in Khairapur of Bardiya district in 1975 captivity on the behavior and survival vary among animal which initiated their conservation eˆorts [3]. �e last species depending on their behavioral •exibility, that is, the ability of animals to modify behavior in adaptive ways over remaining wild population of the species is less than 250 individuals and localized in the small area of the Blackbuck time and space [6, 7]. �rough captive breeding practices, Conservation Area (BCA) in the Bardiya district of the eˆorts are made to breed endangered or threatened animal western lowland, Nepal [4]. In Nepal, the wild habitat of the species so that they could be reintroduced in their natural species is under tremendous pressure from anthropogenic habitats [8]. Captive breeding can play a crucial role in the activities and livestock grazing [2]. �e population of recovery of some species for which eˆective alternatives are Blackbuck in the Blackbuck Conservation Area is unstable unavailable in the short term [9]. �e success of captive and highly con“ned, and therefore, it is susceptible to breeding and reintroduction depends upon a clear under- stochastic deleterious events. In addition to the conservation standing of the reproductive behavior of the species [10]. 2 International Journal of Zoology In ungulates, males and females have contrasting life grassland, 03% riverine forest, 01% mango orchard, and 01% histories, as usually, only the females raise the young, and wetland (site-specific conservation action plan for blackbuck only a handful of territorial males contribute the genetic in Shuklaphanta Wildlife Reserve, Nepal 2016–2020). material [11]. Females become sexually mature at the age of eight months but mate no earlier than two years of age. 2.2. Study Animals. In the beginning of the study period Males mature later, at 1.5 years. Mating takes place (November 2018), the blackbuck enclosure contained a total throughout the year. Gestation is typically six months long, population of 70 animals. (e group was classified into 24 after which a single calf is born. Sexual selection theory adult males, 28 adult females, five subadult males, four predicts that the opportunity for sexual selection is strong subadult females, and nine yearlings. By the end of May when reproductive success varies widely among males, with 2019, the population reached to 86. (e reproductive be- a few males highly successful at mating and many others havior was observed in four adult males only which were in a males failing to mate. An association exists between strong high rut (Figure 2). Each individual was recognized by the sexual selection, high mating polygyny, and high variance in variation in the shape of the horn and the color of the body. male reproductive success [12, 13]. (erefore, it is important (e following morphological features were used to identify to know the environmental and behavioral determinants of the focal male blackbucks among groups [19] (Table 1). successful mating between the male and estrous female. Knowing reproductive behavior is important for establishing a viable population of Blackbuck in Nepal. Male 2.3. Behavioral Observation. (e diurnal activities of blackbucks engage in rutting throughout the year whereas blackbucks were observed daily from 9 : 00 AM to 5 : 00 PM the peak rutting season for females is from July to August for 28 days. (e focus on the reproductive behavior was and from February to March [14]. (e blackbuck territories performed with well-identified animals for an observational have reproductive connotations and are held by adult males shift of three hours (9.00–12.00, 12.00–15.00, and after 15.00 only [15]. During the heat, the blackbucks mark their ter- hours, respectively) by focal sampling method [20]. An ritory by means of urination, defecation, and rubbing ethogram was developed for recording the reproductive preorbital glands against the hard objects. (e scent of the behavior of blackbuck (Table 2). fluid from preorbital glands warns the other males [16]. Excessive fighting also occurs between the males to attract 2.4. Data Analysis. (e association between the behaviors the females during their rutting period [17]. It is important and copulatory mounting was determined by cross-tabu- to understand the reproductive behavior of the endangered lation followed by the Chi-square test. Microsoft Excel was species in order to multiply its population ex-situ. used to analyze the data statistically. Tables were used to A total of 28 blackbucks (22 from Nepalgunj Mini Zoo present the data in simplified form. (e magnitude of the and 6 from Central Zoo, Lalitpur) were reintroduced in effect of factors on behavior for both preceding and suc- Shuklaphanta National Park (SNP) in 2012 as a part of a ceeding the copulatory mounting was measured on the basis national effort to establish a second viable population in of phi value for 2 × 2 cross-tabulation. (e following Nepal (site-specific conservation action plan for blackbuck guidelines were used to determine the magnitude of the in Shuklaphanta National Park, Nepal 2016–2020). Cur- effect size at a 5% level of association of behavioral events rently, 70 individuals of the blackbuck reside in the semi- with the copulatory mounting [21] (Table 3). captive condition of the Hirapurphanta within the SNP [18]. (e success in the re-establishment of the species depends 3. Results on how the animals can cope with the environment and maintain their behavior. Being territorial in nature, the 3.1. Temporal Association of Successful Mounting. reproductive success of blackbuck depends on the adult Blackbucks were in high rut during February–March than breeding dominant males, referred as the alpha males. during July–August. (e higher number of newly born (erefore, we aimed to understand reproductive behavior infants was observed in the month of June/July indicating and major pre and postcopulatory activities of the breeding the higher conception during February–March. A total of males in the semiwild condition of the Hirapurphanta. We 149 copulatory mounting events were documented during employed focal animal sampling methods for behavioral the study period, of which 44.3% were recorded during the sampling and analyzed the association of behavioral states of observational phase of 09.00–12.00 hours, and the least males with successful mounting events. (21.47%) were observed during the afternoon phase (12.00–15.00) (Figure 3). 2. Materials and Methods 2.1. Study Area. (is study was conducted in the blackbuck 3.2. Precopulatory and Courtship Activities. (ere were enclosure of Hirapurphanta in Shuklaphanta National Park various behaviors such as walking, feeding, fighting, chasing, ° ° (80.095–80.361 N, 28.763–29.047 E), Kanchanpur, Nepal resting, and sniffing the vulva of female, showing flehmen (Figure 1). Hirapurphanta has a total area of about 150 response, drinking, urinating, rubbing, and defecating which hectares, but the area of the blackbuck enclosure is 17 preceded copulatory mounting. (ere was a significant as- hectares only. (e blackbuck enclosure lies in the central sociation between walking, feeding, fighting, chasing, rest- part of the Hirapurphanta with 95% of the area covered by ing, and sniffing the vulva of female, showing flehmen International Journal of Zoology 3 0 27,000 54,000 108,000 Meters SNP Hirapur Phanta District Boundary GPS Coordinate Figure 1: Map of Shuklaphanta National Park indicating Hirapurphanta. (a) (b) (c) (d) Figure 2: Adult males (a–d) observed for focal animal sampling. Table 1: Identi“cation features of adult male blackbucks. Animal Number of twists in horn Identi“cation characters A 3 One horn was long and pointed, and the other was small and twisted outward. B 4 Tip of the horns bent inward, long horns, and very broad like a triangle, and body was brown in color. C 4 �ick and parallel horns, most parts of the body were black, and little was brown in color. D 2 Broad and thick horns. One horn was broken during “ghting. Body was black in color. 4 International Journal of Zoology Table 2: Ethogram for monitoring reproductive behaviors of blackbucks inside the captivity of Hirapurphanta. S.N. Behavior Description 1 Sexual display Highly rutted males walked with deliberate steps. Head raised and nose-up and horns consequently laid back. Males marked their territory by rubbing preorbital glands on the ground or by urination and defecation 2 Territorial behavior (depositing their pellets at same place), aggressive calls, “ghting, and chasing other males visiting their territory. Reproductive Sni”ng the vulva of female and showing •ehmen response. Attracting the females through “ghting with other behavior males who visited their territory, showing dominancy, copulatory mounting. 4 Other behaviors Walking, feeding, resting, drinking. Table 3: Phi value and magnitude of eˆect. signi“cant association (p > 0.05) between feeding, “ghting, drinking, urinating, rubbing preorbital glands, and defe- Magnitude of eˆect Phi value (ϕ-value) cating followed after copulatory mounting (Table 5). Strong negative association −1.0 to −0.7 Weak negative association −0.7 to −0.3 4. Discussion Little or no association −0.3 to +0.3 Weak positive association +0.3 to +0.7 4.1. Mating Behavior and Its Seasonality in Blackbuck. Strong positive association +0.7 to +1.0 �e mating system of blackbuck appears to be •exible. �e basic pattern is territorial defense by males [14, 22, 23]. �ey have two peaks in their rutting season, one from March and April, and the other from August to October. During the study period, the blackbucks were found to be in high estrus in the month of March and April. Blackbucks breed throughout the year, but newly born infants were seen more in the month of June. It is well known that environmental conditions may importantly in•uence the reproductive strategies of ungulates. Courtship was initiated by the adult males who searched and followed the female in heat. Fighting occurred when other dominant male visits its territory. Polygynous resource territories, typically held by a single male, generally en- 9.00-12.00 compass important resources such as food, water, or par- 12.00-15.00 ticular types of habitat, and are the most common form of After 15.00 territoriality in polygynous species [24]. During this time, Figure 3: Number of copulatory mountings of blackbuck during the females got attracted by the competitive behavior of the three phases of behavioral sampling. dominant male and chose the strongest male for copulation. �e dominant male in the rutting period sniˆed the vulva of the female, and it gave a head-up with twisting lips. �is response and urinating preceding copulatory mounting, that gesture had been termed as “Flehmen.” During excessive is, p < 0.05 (Table 4). �ere was a strong positive association heat, the males became more excited and chased the females between sni”ng the vulva of female (ϕ-value  0.713) and continuously. During this study, one female died because of showing Flehmen response preceding copulatory mounting forceful mating and “ghting with another dominant male on (ϕ-value  0.799). Little or no association as re•ected by phi 1st June 2019. A dominant male followed two or three fe- value was observed between walking, feeding, “ghting, males and copulated with one which reached the state of chasing, urinating, and resting preceding the copulatory conceiving. mounting. �ere was no signi“cant association between the �e female showed receptivity by waving her tail and behaviors such as rubbing preorbital glands, defecating, and thumping the hind legs on the ground, and the buck fol- drinking preceding copulatory mounting, that is, p > 0.05. lowed the female every time. Mounting was attempted by holding the doe, and the copulation occurred at that stage. After copulation, the doe stood for a while and then started 3.3. Postcopulatory Activities. Among the behavioral events observed succeeding the copulatory mountings, a weak grazing. �e doe sometimes urinated. �e buck took rest due positive association between walking followed after copu- to loss of energy while copulating. Buck sometimes mated latory mounting (ϕ-value  0.347). �ere is a little or no with another doe. It followed the other doe in his territory association between sni”ng the vulva of female and tried to attempt mounting with them. �at was due to (ϕ-value  0.278) and showing •ehmen response the excitement of the copulation. (ϕ-value  0.279) followed after copulatory mounting. Vats and Bhardwaj [25] observed that the strongest Conversely, a weak negative association was recorded be- individual dominated other members of the group. �e tween resting and copulatory mounting. �ere was no encounters happened when one male entered the territory of International Journal of Zoology 5 Table 4: Behavioral events preceding copulatory mounting among the dominant males. Behavioral events Observed frequency Expected frequency X -value p value ϕ-value Walking 47 38.87 3.932 0.047 0.076 Feeding 3 23.7 29.498 <0.0001 −0.210 Fighting 3 11.8 8.802 0.003 −0.0114 Chasing 25 11.5 21.071 <0.0001 0.177 Resting 0 11.8 15.791 <0.0001 −0.153 Sniffing the vulva 57 8.7 341.184 <0.0001 0.713 Flehmen response 69 10.4 428.714 <0.0001 0.799 Drinking 0 2.0 2.290 0.130 −0.058 Urinating 8 3.5 6.917 0.008 0.102 Rubbing preorbital gland 6 3.3 2.747 0.097 0.064 Defecating 0 2.1 2.429 0.119 −0.060 Table 5: Behavioral events succeeding copulatory mounting among the dominant males. Behavioral events Observed frequency Expected frequency X -value p value ϕ-value Walking 28 49.7 31.532 <0.0001 0.347 Feeding 5 7.5 1.905 0.168 −0.085 Fighting 2 3.2 1.015 0.314 −0.062 Chasing 34 23.1 13.199 0.0002 0.224 Resting 2 18.1 35.244 <0.0001 −0.367 Sniffing the vulva 27 15.6 20.250 <0.0001 0.278 Flehmen response 27 15.5 20.440 <0.0001 0.279 Drinking 1 0.5 0.869 0.351 0.057 Urinating 17 15.3 0.430 0.512 0.041 Rubbing preorbital gland 5 5.9 0.307 0.579 −0.034 Defecating 1 2.7 2.315 0.128 −0.094 another male or when the male is in excessive heat. (e buck response, sniffing the vulva, and walking beside the estrous took 5–6 hours in courting the female and finally mounting female were the most frequent behaviors associated with the her copulation terminated courtship. During the copulatory copulatory mounting. (e flehmen or lip-curl behavior is a mount position of the body of the buck, movements of the response of male mammals primarily to female urine, feet are exhibited in perfection to complete the mounting. performed primarily during the reproductive season [30]. It Nutritional requirements vary throughout the repro- is displayed by most ungulate species and is a sexually di- ductive cycle, and strategic feed supplementation can also be morphic behavior seen most frequently when males in- vestigate the genital region of females or freshly voided urine an important tool to improve reproductive efficiency [26]. (e nutritional limitations affect the reproductive process in from females. Functionally, flehmen appears to be related to both genders. (e high energy intake boosts spermatogenesis the transport of nonvolatile chemosensory materials from before fertilization and provides energy reserves to support the oral cavity to the vomeronasal organ [31]. However, the sexual and aggressive behavior during mating [27]. When relative frequency of flehmen in different contexts may not energy intake is maintained and protein intake is reduced, reflect its biological significance [32]. While comparing with there happens a significant reduction in libido and semen reproductive importance, males show higher rates and volume and a trend for reduced sperm output [28]. Male duration of flehmen response to estrous urine than to ungulates incur a substantial energetic cost for engaging in nonestrous urine [33–35] suggesting its importance in the assessment of females’ reproductive condition. reproduction, and they trade off reproductive success with reduced body condition during winter, which may lower (e second most frequent behavioral state associated with the copulatory mounting in blackbuck was sniffing the survival [29]. (e Shuklaphanta National Park used to grow maize and lentils during the dry season. Lentils provide vulva of estrous female. Almost similar observation has been protein which improves the reproductive status whereas described in Reeve’s deer where the highest amount of time maize is necessary because it provides carbohydrates which is invested by the reproductive male in sniffing or licking the provide energy to the blackbuck. Additionally, the blackbucks vulva of female [36]. Olfactory and gustatory stimuli re- are supplemented with wheat, barley, maize, soybean, etc. ceived by the male when the sniffs and licks the vulva of an estrous female evoke sexual excitement expressed by the male’s flehmen response [37]. 4.2. Precopulatory and Postcopulatory Behaviors. Among 11 Females presumably limit the number of mating bouts behavioral states and events observed preceding or suc- with a given male, but the time between successive bouts is ceeding the copulatory mount in blackbuck males, flehmen determined by the dominant male [36]. 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Behavioral Determinants of Mating Success in Blackbuck under Semienclosed Condition: A Cross-Sectional Study in Shuklaphanta National Park, Nepal

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Hindawi International Journal of Zoology Volume 2022, Article ID 8713116, 7 pages https://doi.org/10.1155/2022/8713116 Research Article Behavioral Determinants of Mating Success in Blackbuck under Semienclosed Condition: A Cross-Sectional Study in Shuklaphanta National Park, Nepal 1 1 2 Puja Saud , Juna Neupane , and Mukesh Kumar Chalise Central Department of Zoology, Institute of Science and Technology, Tribhuvan University, Kathmandu, Nepal Nepal Biodiversity Research Society (NEBORS), Lalitpur, Nepal Correspondence should be addressed to Juna Neupane; zunaneupane@gmail.com Received 1 April 2022; Revised 19 June 2022; Accepted 12 July 2022; Published 5 August 2022 Academic Editor: Irene Pellegrino Copyright © 2022 Puja Saud et al. �is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Knowing reproductive behavior is important for establishing a viable population of nationally endangered blackbuck in Nepal. �is study explored the behavioral events and states associated with copulatory mountings in blackbuck by focal animal sampling method under semienclosed conditions of Hirapurphanta within Shuklaphanta National Park, Nepal. We recorded diˆerent behavioral initiations for the mating reproductive activities and the behaviors followed after copulatory mounting. Four dominant adult males were used as a subject. �e highest number of copulatory mountings was observed during the months of Febru- ary–March. Among the diurnal observational phases, the highest number of copulatory mountings was recorded during 9.00–12.00 hours followed by 15.00 hours onwards. Copulatory mounting was signi“cantly preceded by sni”ng the vulva of female and showing •ehmen response, whereas copulatory mounting was followed by walking. eˆorts in the wild condition, ex-situ breeding in the captivity 1. Introduction and introduction to the semiwild habitats are undergoing. Blackbucks (Antilope cervicapra, Bovidae, Artiodactyla) Captive breeding is one of the best ways to ensure the were formerly widespread across the Indian subcontinent future existence of any plant or animal. However, the captive but are now limited to only small, scattered herds in Nepal, environment can be aˆecting the animal behavior mainly in India, and Pakistan due to excessive hunting and reduced four ways: the regular presence of large numbers of unfa- habitat range [1]. Blackbuck is one of the highly threatened miliar and behaviorally diverse (passive-active) humans, and nationally protected species in Nepal [2]. �ey were restricted and unchanging space, forced proximity to other thought almost extinct until a small herd of nine individuals animal species, and human management [5]. Eˆects of was discovered in Khairapur of Bardiya district in 1975 captivity on the behavior and survival vary among animal which initiated their conservation eˆorts [3]. �e last species depending on their behavioral •exibility, that is, the ability of animals to modify behavior in adaptive ways over remaining wild population of the species is less than 250 individuals and localized in the small area of the Blackbuck time and space [6, 7]. �rough captive breeding practices, Conservation Area (BCA) in the Bardiya district of the eˆorts are made to breed endangered or threatened animal western lowland, Nepal [4]. In Nepal, the wild habitat of the species so that they could be reintroduced in their natural species is under tremendous pressure from anthropogenic habitats [8]. Captive breeding can play a crucial role in the activities and livestock grazing [2]. �e population of recovery of some species for which eˆective alternatives are Blackbuck in the Blackbuck Conservation Area is unstable unavailable in the short term [9]. �e success of captive and highly con“ned, and therefore, it is susceptible to breeding and reintroduction depends upon a clear under- stochastic deleterious events. In addition to the conservation standing of the reproductive behavior of the species [10]. 2 International Journal of Zoology In ungulates, males and females have contrasting life grassland, 03% riverine forest, 01% mango orchard, and 01% histories, as usually, only the females raise the young, and wetland (site-specific conservation action plan for blackbuck only a handful of territorial males contribute the genetic in Shuklaphanta Wildlife Reserve, Nepal 2016–2020). material [11]. Females become sexually mature at the age of eight months but mate no earlier than two years of age. 2.2. Study Animals. In the beginning of the study period Males mature later, at 1.5 years. Mating takes place (November 2018), the blackbuck enclosure contained a total throughout the year. Gestation is typically six months long, population of 70 animals. (e group was classified into 24 after which a single calf is born. Sexual selection theory adult males, 28 adult females, five subadult males, four predicts that the opportunity for sexual selection is strong subadult females, and nine yearlings. By the end of May when reproductive success varies widely among males, with 2019, the population reached to 86. (e reproductive be- a few males highly successful at mating and many others havior was observed in four adult males only which were in a males failing to mate. An association exists between strong high rut (Figure 2). Each individual was recognized by the sexual selection, high mating polygyny, and high variance in variation in the shape of the horn and the color of the body. male reproductive success [12, 13]. (erefore, it is important (e following morphological features were used to identify to know the environmental and behavioral determinants of the focal male blackbucks among groups [19] (Table 1). successful mating between the male and estrous female. Knowing reproductive behavior is important for establishing a viable population of Blackbuck in Nepal. Male 2.3. Behavioral Observation. (e diurnal activities of blackbucks engage in rutting throughout the year whereas blackbucks were observed daily from 9 : 00 AM to 5 : 00 PM the peak rutting season for females is from July to August for 28 days. (e focus on the reproductive behavior was and from February to March [14]. (e blackbuck territories performed with well-identified animals for an observational have reproductive connotations and are held by adult males shift of three hours (9.00–12.00, 12.00–15.00, and after 15.00 only [15]. During the heat, the blackbucks mark their ter- hours, respectively) by focal sampling method [20]. An ritory by means of urination, defecation, and rubbing ethogram was developed for recording the reproductive preorbital glands against the hard objects. (e scent of the behavior of blackbuck (Table 2). fluid from preorbital glands warns the other males [16]. Excessive fighting also occurs between the males to attract 2.4. Data Analysis. (e association between the behaviors the females during their rutting period [17]. It is important and copulatory mounting was determined by cross-tabu- to understand the reproductive behavior of the endangered lation followed by the Chi-square test. Microsoft Excel was species in order to multiply its population ex-situ. used to analyze the data statistically. Tables were used to A total of 28 blackbucks (22 from Nepalgunj Mini Zoo present the data in simplified form. (e magnitude of the and 6 from Central Zoo, Lalitpur) were reintroduced in effect of factors on behavior for both preceding and suc- Shuklaphanta National Park (SNP) in 2012 as a part of a ceeding the copulatory mounting was measured on the basis national effort to establish a second viable population in of phi value for 2 × 2 cross-tabulation. (e following Nepal (site-specific conservation action plan for blackbuck guidelines were used to determine the magnitude of the in Shuklaphanta National Park, Nepal 2016–2020). Cur- effect size at a 5% level of association of behavioral events rently, 70 individuals of the blackbuck reside in the semi- with the copulatory mounting [21] (Table 3). captive condition of the Hirapurphanta within the SNP [18]. (e success in the re-establishment of the species depends 3. Results on how the animals can cope with the environment and maintain their behavior. Being territorial in nature, the 3.1. Temporal Association of Successful Mounting. reproductive success of blackbuck depends on the adult Blackbucks were in high rut during February–March than breeding dominant males, referred as the alpha males. during July–August. (e higher number of newly born (erefore, we aimed to understand reproductive behavior infants was observed in the month of June/July indicating and major pre and postcopulatory activities of the breeding the higher conception during February–March. A total of males in the semiwild condition of the Hirapurphanta. We 149 copulatory mounting events were documented during employed focal animal sampling methods for behavioral the study period, of which 44.3% were recorded during the sampling and analyzed the association of behavioral states of observational phase of 09.00–12.00 hours, and the least males with successful mounting events. (21.47%) were observed during the afternoon phase (12.00–15.00) (Figure 3). 2. Materials and Methods 2.1. Study Area. (is study was conducted in the blackbuck 3.2. Precopulatory and Courtship Activities. (ere were enclosure of Hirapurphanta in Shuklaphanta National Park various behaviors such as walking, feeding, fighting, chasing, ° ° (80.095–80.361 N, 28.763–29.047 E), Kanchanpur, Nepal resting, and sniffing the vulva of female, showing flehmen (Figure 1). Hirapurphanta has a total area of about 150 response, drinking, urinating, rubbing, and defecating which hectares, but the area of the blackbuck enclosure is 17 preceded copulatory mounting. (ere was a significant as- hectares only. (e blackbuck enclosure lies in the central sociation between walking, feeding, fighting, chasing, rest- part of the Hirapurphanta with 95% of the area covered by ing, and sniffing the vulva of female, showing flehmen International Journal of Zoology 3 0 27,000 54,000 108,000 Meters SNP Hirapur Phanta District Boundary GPS Coordinate Figure 1: Map of Shuklaphanta National Park indicating Hirapurphanta. (a) (b) (c) (d) Figure 2: Adult males (a–d) observed for focal animal sampling. Table 1: Identi“cation features of adult male blackbucks. Animal Number of twists in horn Identi“cation characters A 3 One horn was long and pointed, and the other was small and twisted outward. B 4 Tip of the horns bent inward, long horns, and very broad like a triangle, and body was brown in color. C 4 �ick and parallel horns, most parts of the body were black, and little was brown in color. D 2 Broad and thick horns. One horn was broken during “ghting. Body was black in color. 4 International Journal of Zoology Table 2: Ethogram for monitoring reproductive behaviors of blackbucks inside the captivity of Hirapurphanta. S.N. Behavior Description 1 Sexual display Highly rutted males walked with deliberate steps. Head raised and nose-up and horns consequently laid back. Males marked their territory by rubbing preorbital glands on the ground or by urination and defecation 2 Territorial behavior (depositing their pellets at same place), aggressive calls, “ghting, and chasing other males visiting their territory. Reproductive Sni”ng the vulva of female and showing •ehmen response. Attracting the females through “ghting with other behavior males who visited their territory, showing dominancy, copulatory mounting. 4 Other behaviors Walking, feeding, resting, drinking. Table 3: Phi value and magnitude of eˆect. signi“cant association (p > 0.05) between feeding, “ghting, drinking, urinating, rubbing preorbital glands, and defe- Magnitude of eˆect Phi value (ϕ-value) cating followed after copulatory mounting (Table 5). Strong negative association −1.0 to −0.7 Weak negative association −0.7 to −0.3 4. Discussion Little or no association −0.3 to +0.3 Weak positive association +0.3 to +0.7 4.1. Mating Behavior and Its Seasonality in Blackbuck. Strong positive association +0.7 to +1.0 �e mating system of blackbuck appears to be •exible. �e basic pattern is territorial defense by males [14, 22, 23]. �ey have two peaks in their rutting season, one from March and April, and the other from August to October. During the study period, the blackbucks were found to be in high estrus in the month of March and April. Blackbucks breed throughout the year, but newly born infants were seen more in the month of June. It is well known that environmental conditions may importantly in•uence the reproductive strategies of ungulates. Courtship was initiated by the adult males who searched and followed the female in heat. Fighting occurred when other dominant male visits its territory. Polygynous resource territories, typically held by a single male, generally en- 9.00-12.00 compass important resources such as food, water, or par- 12.00-15.00 ticular types of habitat, and are the most common form of After 15.00 territoriality in polygynous species [24]. During this time, Figure 3: Number of copulatory mountings of blackbuck during the females got attracted by the competitive behavior of the three phases of behavioral sampling. dominant male and chose the strongest male for copulation. �e dominant male in the rutting period sniˆed the vulva of the female, and it gave a head-up with twisting lips. �is response and urinating preceding copulatory mounting, that gesture had been termed as “Flehmen.” During excessive is, p < 0.05 (Table 4). �ere was a strong positive association heat, the males became more excited and chased the females between sni”ng the vulva of female (ϕ-value  0.713) and continuously. During this study, one female died because of showing Flehmen response preceding copulatory mounting forceful mating and “ghting with another dominant male on (ϕ-value  0.799). Little or no association as re•ected by phi 1st June 2019. A dominant male followed two or three fe- value was observed between walking, feeding, “ghting, males and copulated with one which reached the state of chasing, urinating, and resting preceding the copulatory conceiving. mounting. �ere was no signi“cant association between the �e female showed receptivity by waving her tail and behaviors such as rubbing preorbital glands, defecating, and thumping the hind legs on the ground, and the buck fol- drinking preceding copulatory mounting, that is, p > 0.05. lowed the female every time. Mounting was attempted by holding the doe, and the copulation occurred at that stage. After copulation, the doe stood for a while and then started 3.3. Postcopulatory Activities. Among the behavioral events observed succeeding the copulatory mountings, a weak grazing. �e doe sometimes urinated. �e buck took rest due positive association between walking followed after copu- to loss of energy while copulating. Buck sometimes mated latory mounting (ϕ-value  0.347). �ere is a little or no with another doe. It followed the other doe in his territory association between sni”ng the vulva of female and tried to attempt mounting with them. �at was due to (ϕ-value  0.278) and showing •ehmen response the excitement of the copulation. (ϕ-value  0.279) followed after copulatory mounting. Vats and Bhardwaj [25] observed that the strongest Conversely, a weak negative association was recorded be- individual dominated other members of the group. �e tween resting and copulatory mounting. �ere was no encounters happened when one male entered the territory of International Journal of Zoology 5 Table 4: Behavioral events preceding copulatory mounting among the dominant males. Behavioral events Observed frequency Expected frequency X -value p value ϕ-value Walking 47 38.87 3.932 0.047 0.076 Feeding 3 23.7 29.498 <0.0001 −0.210 Fighting 3 11.8 8.802 0.003 −0.0114 Chasing 25 11.5 21.071 <0.0001 0.177 Resting 0 11.8 15.791 <0.0001 −0.153 Sniffing the vulva 57 8.7 341.184 <0.0001 0.713 Flehmen response 69 10.4 428.714 <0.0001 0.799 Drinking 0 2.0 2.290 0.130 −0.058 Urinating 8 3.5 6.917 0.008 0.102 Rubbing preorbital gland 6 3.3 2.747 0.097 0.064 Defecating 0 2.1 2.429 0.119 −0.060 Table 5: Behavioral events succeeding copulatory mounting among the dominant males. Behavioral events Observed frequency Expected frequency X -value p value ϕ-value Walking 28 49.7 31.532 <0.0001 0.347 Feeding 5 7.5 1.905 0.168 −0.085 Fighting 2 3.2 1.015 0.314 −0.062 Chasing 34 23.1 13.199 0.0002 0.224 Resting 2 18.1 35.244 <0.0001 −0.367 Sniffing the vulva 27 15.6 20.250 <0.0001 0.278 Flehmen response 27 15.5 20.440 <0.0001 0.279 Drinking 1 0.5 0.869 0.351 0.057 Urinating 17 15.3 0.430 0.512 0.041 Rubbing preorbital gland 5 5.9 0.307 0.579 −0.034 Defecating 1 2.7 2.315 0.128 −0.094 another male or when the male is in excessive heat. (e buck response, sniffing the vulva, and walking beside the estrous took 5–6 hours in courting the female and finally mounting female were the most frequent behaviors associated with the her copulation terminated courtship. During the copulatory copulatory mounting. (e flehmen or lip-curl behavior is a mount position of the body of the buck, movements of the response of male mammals primarily to female urine, feet are exhibited in perfection to complete the mounting. performed primarily during the reproductive season [30]. It Nutritional requirements vary throughout the repro- is displayed by most ungulate species and is a sexually di- ductive cycle, and strategic feed supplementation can also be morphic behavior seen most frequently when males in- vestigate the genital region of females or freshly voided urine an important tool to improve reproductive efficiency [26]. (e nutritional limitations affect the reproductive process in from females. Functionally, flehmen appears to be related to both genders. (e high energy intake boosts spermatogenesis the transport of nonvolatile chemosensory materials from before fertilization and provides energy reserves to support the oral cavity to the vomeronasal organ [31]. However, the sexual and aggressive behavior during mating [27]. When relative frequency of flehmen in different contexts may not energy intake is maintained and protein intake is reduced, reflect its biological significance [32]. While comparing with there happens a significant reduction in libido and semen reproductive importance, males show higher rates and volume and a trend for reduced sperm output [28]. Male duration of flehmen response to estrous urine than to ungulates incur a substantial energetic cost for engaging in nonestrous urine [33–35] suggesting its importance in the assessment of females’ reproductive condition. reproduction, and they trade off reproductive success with reduced body condition during winter, which may lower (e second most frequent behavioral state associated with the copulatory mounting in blackbuck was sniffing the survival [29]. (e Shuklaphanta National Park used to grow maize and lentils during the dry season. Lentils provide vulva of estrous female. Almost similar observation has been protein which improves the reproductive status whereas described in Reeve’s deer where the highest amount of time maize is necessary because it provides carbohydrates which is invested by the reproductive male in sniffing or licking the provide energy to the blackbuck. Additionally, the blackbucks vulva of female [36]. Olfactory and gustatory stimuli re- are supplemented with wheat, barley, maize, soybean, etc. ceived by the male when the sniffs and licks the vulva of an estrous female evoke sexual excitement expressed by the male’s flehmen response [37]. 4.2. Precopulatory and Postcopulatory Behaviors. Among 11 Females presumably limit the number of mating bouts behavioral states and events observed preceding or suc- with a given male, but the time between successive bouts is ceeding the copulatory mount in blackbuck males, flehmen determined by the dominant male [36]. 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Journal

International Journal of ZoologyHindawi Publishing Corporation

Published: Aug 5, 2022

References