Ecological Determinants of Chital (Axis axis) Reproductive Seasonality in Kanha Tiger Reserve

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While tropical and equatorial regions exhibit continual breeding due to stable resources, tropical India experiences pronounced seasonality in forage availability due to monsoonal rainfall. This study investigates the breeding timing of chital ( Axis axis ) in the tropical deciduous forests of Central India at Kanha Tiger Reserve, where forage availability varies seasonally. Field observations focused on antler conditions in males and lactation signs in females to estimate breeding timing. Rutting peak, as determined by males with hard antlers, coincided with nutritional peak induced by monsoon, which probably was the cue for ovulation in females. Lactation, the most nutritionally demanding period, peaked with the post-fire sprouting of herbaceous vegetation in April and fawn weaning synchronized with ample food availability. This synchrony between reproductive cycles and forage availability emphasizes the adaptive strategies of chital to maximize offspring survival. Furthermore, the peak in tiger births (54%) coincided with the fawning peak of chital. making them vulnerable prey during a nutritionally demanding period for tigresses, who are restricted in their movements near vulnerable cubs. Understanding the links between trophic levels and their seasonality offers important insights for wildlife management and conservation strategies. Biological sciences/Ecology Biological sciences/Zoology Earth and environmental sciences/Ecology Earth and environmental sciences/Environmental sciences Fawning Lactation Rutting Nutrition Tiger Births Figures Figure 1 Figure 2 Introduction Synchrony or birthing peaks are seen as a response to either seasonal environments or as an anti-predatory adaptation (Roeder et al. 2022 ). Tropical and equatorial mammals exhibit a lack of seasonality in breeding due to uniform resource availability. However, tropical India is highly seasonal in resource availability due to the rainfall being primarily restricted to the monsoon season. Sinclair et al. ( 2000 ) studied 13 African ungulates and found that their breeding synchronizes with seasons of abundant forage resources, an adaptation that enhances offspring survival by ensuring that young are born during favourable conditions. Rainfall patterns also significantly influence ungulate reproductive behaviour; Priyadarshini ( 2005 ) notes that rainfall affects food availability, which in turn determines reproductive success. Increased rainfall can lead to lush vegetation, providing better nutrition for pregnant or lactating females and improving offspring survival rates. The reproductive patterns of ungulates can vary significantly based on environmental conditions. Ungulates are timed to coincide with their breeding season with periods of highest plant quality to support offspring nutrition (Mduma et al. 1999 ). This adaptation ensures that lactating females and weaned offspring obtain the required energy and protein from the forage (Western 1979 ; Bronson 1989 ; Owen-Smith et al. 2005 ; Priyadarshini 2005 ). These relationships underscore the importance of understanding ecological dynamics to better manage ungulate populations and their habitats. As environmental conditions continue to change due to human impact, ongoing research will be required to understand how ungulates adapt to these challenges. Changes in forage quantity and quality significantly impact the body condition of ungulates, especially females during gestation and lactation, when their energy and protein requirements are heightened. Studies by Oftedal 1985 ; Robbins 1993 ; Parker et al. 1999 ; Gaillard et al. 2000 ; Sinclair et al. 2000 ; and Cook et al. 2001 all emphasize the critical role that adequate nutrition plays in ensuring the health and reproductive success of these animals during these demanding periods. The onset of oestrus in females is significantly affected by conditions required for a successful pregnancy and subsequent lactation, such as food availability or favourable weather conditions (McGinnes & Downing 1977 ; Asher 2011 ), as increased time and energy expenditures for food searching can lead to higher mortality in reproducing adults (Robbins 1993 ). We hypothesize that after a stressful dry summer, the new flush in vegetation caused by the monsoon rains should be the cue for ovulation in female ungulates, including chital in India. In most of the cervids, antler cycles serve as external indicators of internal reproductive changes (Bubenik et al. 2002 ). In most deer species, males undergo a yearly cycle characterized by the growth and subsequent shedding of antlers (Bubenik et al. 1987 ; Tomás 1995 ; Ungerfeld et al. 2008 ). Antlers play a crucial role in male-male confrontations to establish dominance and secure mates, serving as an indication to females of male fitness and condition during the rutting season, which is a period of intense mating activity (Bubenik et al. 1991 ; Clements et al. 2010 ; Vanpé et al. 2010 ; Heckeberg 2017 ). Testosterone levels peak seasonally during the rut season (Bubenik 1990 ), and when testosterone levels fall below threshold after the rutting season, antlers are cast off in most cervids (Putman 1988 ; Bartoš 1990 ; Bubenik 1990 ). Consequently, peak rutting activity often coincides with the period when the largest proportion of males exhibit burnished antlers, indicative of their reproductive readiness (Raman 1998 ; Ramesh et al. 2013 ; Bhusal et al. 2020 ). In deer, observing actual fawning is a rare event and is often studied by implanting radio transmitters in the vagina of pregnant females, which are expelled out at the time of parturition (Bowman & Jacobson 1998 ; Conant et al. 2020 ). This allows for the study of timing and specific location of parturition. We indexed the timing of fawning by estimating the proportion of lactating females in the population (Priyadarshini 2005 ; Jhala & Isvaran 2016 ). Many studies on breeding seasonality have been conducted on North American and African ungulates, but in the Indian subcontinent only a few ungulate species have been studied for their reproductive patterns (Rice 1988 ; Raman 1998 ). However, these studies have primarily focused on observing the seasonality of the antler cycle. Management decisions like prey augmentation (Phumanee et al. 2021 ), where animals from high-density areas were captured and moved to low-density areas to increase the carrying capacity of large carnivores, also need information about rutting and fawning. Translocation during prey augmentation in the fawning season can cause stress due to possible mother-fawn separation, and aggressive behaviour of males that are in hard antlers during the rutting season can increase mortality, making it essential to understand the breeding season of the source chital population beforehand. In this study, we record the reproductive condition of chital throughout the year from representative samples to decipher seasonality and its likely drivers in the dry deciduous forest system of Kanha National Park. Methods Study area Kanha Tiger Reserve is located in the Satpura Maikal hill ranges of the Central Indian highlands (Rodgers & Panwar 1988 ). It was among the first nine Tiger Reserves gazetted in 1973. It is located between 80°26’10’’ E to 81°04’40.00’’E and 22°01’5.0” N to 27°27’48.00” N in Balghat and Mandla districts of Madhya Pradesh. The area of the Core Zone is 917.43 km2, and that of the Buffer Zone is 1134.39 km 2 , adding up to 2051.82 km 2 as the total area of Kanha Tiger Reserve. The Tiger Reserve has flat hilltops, varying degrees of slopes, and meadows in the valleys, which offer unique, diverse types of wildlife habitat, forming ideal niches for various species of plants and animals. There are three distinct seasons in Kanha Tiger Reserve. These are monsoon, summer, and winter. Summer extends from mid-February to June end, monsoon is from June end week to mid-October, and winter is from mid-October to mid-February. Due to a great variation in temperature, humidity, wind velocity, and precipitation in different seasons, these factors serve as regulators of vegetation and the habits of wild animals in the reserve. Kanha Tiger Reserve lies in a dry deciduous zone, and the availability of water is a very important factor for the survival of animals and plants. Major forest types of the area can be divided into four categories: sal forest, miscellaneous forest, bamboo mixed forest, and grasslands. Controlled burning practices during February are carried out in the grasslands of Kanha each year for habitat improvement and management purposes. Field methods Estimating proportion of population in breeding condition Observation of actual mating events and fawning in wild populations is rare, especially in tropical forest systems. We therefore indexed breeding condition in males by assessing their antler condition (Bubenik et al. 2002 ; Ramesh et al. 2013 ) and fawning in females by assessing their lactation state (Fryxell 1987 ; Eberhardt et al. 1996 ; Laurian et al. 2000 ; Priyadarshini 2005 ). For obtaining an unbiased representation of the population in samples, we conducted intensive surveys along fixed vehicle routes to observe the breeding status of male and female chital groups. Each route was visited once a week. Initially, the population was categorized into adults, sub-adults, and juveniles, following Schaller ( 1967 ). For assessing breeding signs, only adult individuals were considered. To assess the involvement of sub-adult females in breeding, a total of 200 sub-adult females were observed for lactation signs during the peak lactation period. We subdivided adult males into hard antler, velvet, and shed antler male groups as per their antler stages at different times of the year (Figure S1 ). We expect that the peak breeding season will coincide with the time when the largest proportion of bucks exhibit hard antlers. To determine the fawning season, lactation signs were recorded whenever an adult female was observed, as the presence of lactating mothers indicated the birth of a fawn (Fryxell 1987 ; Eberhardt et al. 1996 ; Laurian et al. 2000 ; Priyadarshini 2005 ). In chital udders and teats become visible only a few days before parturition and remain prominent up to four months after parturition (Weigl 2005 ; Myhrvold et al. 2015 ) (Figure S2). Analytical method Mating and fawning season Data on various stages of male antlers and lactating females were pooled month-wise for exploratory analysis. Then the data were categorized into three distinct seasons: winter, summer, and monsoon, and reanalysed. Since the same route was followed weekly for observations, the computation was done using multiple sampling with replacement approach (Skalski et al. 2010 ). NDVI change Chital are the primary grazers at Kanha Tiger Reserve as more than 50% of their diet consists of grasses (Awasthi 2020 ). So change in the condition of grass affects the body requirements of chital. To see the monthly productivity in the grasslands of Kanha, we analysed the Normalized Difference Vegetation Index (NDVI) of the grasslands of Kanha for the past 10 years from satellite imagery of LANDSAT 8 (USGS) in Google Earth Engine (Gorelick et al. 2017 ). A harmonic model for monthly change in NDVI was fitted to the data to see monthly changes in the grasslands of Kanha (Fig S3). When sprouting begins in grasslands, the NDVI value starts increasing from a low as the grasses turn green replacing the dry grass or bare land left after burning. Then monthly average values of NDVI were plotted to evaluate the synchrony between grass sprouting season and chital breeding in Kanha. Birth of tiger cubs We used information from Kumar ( 2019 ) on the number of litters born each month for tigers in Kanha. Kumar ( 2019 ) determined the birth month of tiger litters by aging the cubs. The birth records and the month of parturition were obtained through continuous sightings by experienced forest staff, researchers, camera trap pictures, and handheld photographs taken by tourists who shared the date and location information. Information on 31 litter births was recorded with reasonable certainty. Once the month of birth was ascertained, the relationship between chital fawning and tiger cub was explored by looking at the time of birth of offspring of both prey and predator. Results Fawning occurred throughout the year as lactation was recorded in all months. However, the proportion of lactating females in the population peaked in March-April. April has the highest proportion of lactating females (78 ± 0.8%), followed by March (73.5 ± 1.1%), while November recorded the lowest proportion (6.93 ± 0.37%) (Fig. 1 , Table S1 ). Seasonally, summer had the highest proportion of lactating females (72.26 ± 0.24%), followed by monsoon (48.9 ± 0.46%), and winter (30.79 ± 0.34%) (Fig. 1 , Table S1 ). None of the 200 sampled sub-adult females were observed to be lactating. In July, nearly all adult males have hard antlers (97.05 ± 0.15%). Between May and November, over 60% of the male population displayed hard antlers, while more than 50% of the adult male population sprouted velvet-covered antlers between December and April (Fig. 1 , table S2). The proportion of shed antler male observation is low due to the short interval between antler shedding and the emergence of new antlers. In Kanha Tiger Reserve, grassland growth peaks during the monsoon and post-monsoon periods but declines during the dry winter season with minimal rainfall. The NDVI value during the post-fire season (April) shows an upward trend indicating the emergence of freshly grown grass shoots. Lactating peaked at the time of the new grass flush when the NDVI values start having an upward trend (Fig. 1 , Table S3) after a lowest in March. Births of tiger litters were observed throughout the year with a peak in December. Their weaning started after 3–4 months which coincided with the same time of fawning (Fig. 2 ). Discussion There seems to be a debate regarding the existence of distinct seasonality in chital, particularly in tropical regions where food and water are relatively stable throughout the year. Krishnan ( 1972 ) states that chital lacks any birth peak throughout peninsular India, but many studies found distinct seasonal peaks in chital rutting behaviour in India (Table S4). Studies were done in Hawaii by Graff & Nichols ( 1966 ), Texas by Ables ( 1977 ), Australia by Kelly et al. ( 2022 ), Nepal by Bhusal et al. ( 2020 ), and Mishra ( 1982 ) also found a distinct peak in rutting behaviour of chital depending on environmental factors like day length and resource availability. The Central Indian landscape has distinct seasonality, which affects the availability and quality of forage during different times of the year. Mixed foragers like Chital shift their diet towards browsing during summer when grass nutrient quality declines (Awasthi 2020 ; Watter et al. 2020 ), emphasizing the importance of dietary quality. The dry grasses become non-nutritious and are filled with silica and lignin content (Kayongo-Male et al. 1976 ). Due to the burning of grasslands during February to reduce the fuel load of grasses, preventing uncontrolled fire during the summer season, fresh biomass appears post-fire season, which is high in crude protein (White 1983 ; Akbarinia & Koocheki 1992 ; Arzani et al. 1998 , 2004 ) and other minerals like calcium and magnesium (Devi et al. 2022 ). We found that the breeding peak of chital is synchronous with the appearance of fresh grass forage biomass, i.e., the months of March and April. Moreover, fawns born at this time were weaned off during the monsoon period, when there is ample forage available for them to feed on. Ogutu et al. ( 2010 ) noticed the birth peak of ungulates coincided with seasonal rainfall and the appearance of green foliage in tropical and subtropical regions. Clutton-Brock et al. ( 1989 ) also state that ungulate breeding is affected by food availability, and this hypothesis is also supported by Ahrestani et al. ( 2012 ), who conclude that chital also follows this pattern. In Kanha, rutting occurs in late summer and monsoon season, although some hard antler males are present in the population throughout the year. Deer females conceive only when they have good body condition (Flajšman et al. 2017 ; Paoli et al. 2018 ), and Kelly et al. ( 2022 ) found that this is true for chitals also. In Kanha, chital has better body condition when ample food is available (Awasthi 2020 ). Similarly, fawning starts in January and reaches a peak during April. Since fawning is represented by lactating females and lactation occurs up to 4 months in chitals, it is difficult to determine the exact fawning peak. Peak in tiger cub birth is the same time when more than half of the female chital were lactating. When weaning of tiger cubs starts after a few months, it is the same time when the number of lactating females is also highest. When weaning starts, the need for prey also increases, as the mother needs to provide food for weaned cubs. During this period, chitals are most vulnerable to predation, with males being in hard antlers and preoccupied with the rut, females caring for fawns and weak due to lactation, and ample availability of fawns. Tigers in the Kanha landscape showed a functional response by selective predation on the abundant chital fawns and rutting males (Jhala et al. 2014 ), resulting in numerical response by timing their birth peak at this time. Similar pattern has also been reported in mountain lions ( Puma concolor ) by Jansen & Jenks ( 2012 ). The mountain lion birth peak is synchronized with the birth of North American ungulates in May-June. There are certain cues like resource abundance that track ecological rhythms that influence reproductive behaviour. The intricate relationship between seasonal forage dynamics, reproductive patterns, nutritional ecology, and predation underscores the importance of understanding these dynamics for effective wildlife management and conservation planning. Birthing and rutting peaks of chital in Kanha, which is a deciduous forest-grassland system, were determined by resource availability and were unlikely to be in response to predation. Conclusion This study on the seasonality of chital, particularly in tropical regions with stable food and water availability, highlights the complexity of their reproductive behaviour. In the central Indian landscape, seasonality significantly affects forage availability and quality. The management practices in tiger reserves, such as controlled burning, enhance forage quality by promoting the growth of nutrient-rich fresh biomass. This study finds that the breeding peak of chital in Kanha synchronizes with the appearance of fresh grass forage in March and April after controlled burning, providing crucial nutrition to lactating females. Rutting occurs in late summer and monsoon, while fawning peaks in April, which represents a proxy for the presence of lactating females, making them and their fawns more vulnerable to predation. The alignment of tiger reproductive biology with the peak availability of chital fawns suggests a functional predator response. The interplay between seasonal forage dynamics, reproductive patterns, nutritional ecology, and predation is vital for effective wildlife management and conservation. Understanding these dynamics in the deciduous forest-grassland system of Kanha reveals that chital birthing and rutting peaks are driven more by resource availability than predation pressures. Effective conservation strategies must consider these intricate ecological relationships to support the sustainable management of chital populations and their habitats. Declarations Acknowledgement We thank Chief Wildlife Warden of Madhya Pradesh and management of Kanha Tiger Reserve for permissions and logistics for the study. We thank our field assistants Nirottam and Kanhaiya and the team of forest guards for their help in field data collection. We thank field director Kanha, S.K. Singh for logistic support . This study was funded by National Tiger Conservation Authority, Ministry of Environment, Forest and Climate change, Government of India. Authors' contributions: YVJ & QQ conceived, supervised, procured the resources for the study, SG did the field data collection, SG & YVJ did the data analysis, SG, UK and YVJ wrote the MS, and all the authors approved the MS. Availability of data and material: All data generated and analysed during this study are included in this published article and its supplementary information files. Ethics approval: The study was based on observation. All observation was made in the animal’s natural habitat without any physical interference. The study complied with the ethical guidelines issued by Chief Wildlife Warden through permit No. Serial no./D.M.II/Research-213/9027. Competing interests: The authors declare that they have no conflict of interest. References Ables, E.D. (1977). The axis deer in Texas Vol. 2. College Station: Texas A&M University Press. 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Ramesh, T., Kalle, R., Sankar, K., Qureshi, Q., & Downs, C.T. (2013). Aspects of breeding biology of chital ( Axis axis ) and sambar ( Rusa unicolor ) in the Western Ghats. Acta Ethologica 16: 147–155. Rice, C.G. (1988). Reproductive biology of Nilgiri tahr, Hemitragus hylocrius (Mammalia: Bovidae. Journal of Zoology 214(2): 269–284. Robbins, C.T. (1993). Wildlife feeding and nutrition . Academic Press, New York. Rodgers, W. & Panwar, H. (1988). Planning a wildlife protected area network in India Roeder, D.V., Husak, M.S., Murphy, M.T., & Patten, M.A. (2022). Combined roles for breeding synchrony, habitat and scale as predictors of extrapair paternity. Animal Behaviour 194: 139–150.; https://doi.org/10.1016/j.anbehav.2022.09.016 Schaller, G.B. (1967). The deer and the tiger a study of wildlife in India . University of Chicago Press. Sinclair, A.R.E., Mduma, S.A.R., & Arcese, P. (2000). What determines phenology and synchrony of ungulate breeding in the Serengeti? Ecology 81: 2100–2111. Skalski, J.R., Ryding, K.E., & Millspaugh, J.J. (2010). Wildlife demography: analysis of sex, age, and count data . Elsevier Academic Press, Burlington, MA. Tomás, W.M. (1995). Seasonality of the antler cycle of pampas deer ( Ozotoceros bezoarticus leucogaster ) from the Pantanal Wetland, Brazil. Studies on Neotropical Fauna and Environment 30(4): 221–227. Ungerfeld, R., González-Sierra, U.T., & Bielli, A. (2008). Seasonal antler cycle in a herd of pampas deer ( Ozotoceros bezoarticus ) in Uruguay. Mammalian Biology 73(5): 388–391. Vanpé, C., Gaillard, J.M., Kjellander, P., Liberg, O., Delorme, D., & Hewison, A.M. (2010). Assessing the intensity of sexual selection on male body mass and antler length in roe deer Capreolus capreolus : is bigger better in a weakly dimorphic species? Oikos 119(9): 1484–1492. Watter, K., Baxter, G., Brennan, M., Pople, T., & Murray, P. (2020). Seasonal diet preferences of chital deer in the northern Queensland dry tropics, Australia. The Rangeland Journal 42(3): 211.; https://doi.org/10.1071/RJ20015 Weigl, R. (2005). Longevity of mammals in captivity: from the living collections of the world ; a list of mammalian longevity in captivity . Schweizerbart, Stuttgart, 214pp. Western, D. (1979). Size, life history and ecology in mammals. African Journal of Ecology 17(4): 185–204. White, L.M. (1983). Seasonal changes in yield, digestibility, and crude protein of vegetative and floral tillers of two grasses. Journal of Range Management 36: 402–404. Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-5756049","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":400510523,"identity":"5b3f9be5-913c-43fc-bcc0-4a6026ad24ec","order_by":0,"name":"Shravana Goswami","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAElEQVRIiWNgGAWjYDACCTSuHIg88IAULcZgLQkkaGFIbACR+LTIz+59+Jmn5o68vHv7M4mfORbp88MOPwTaYien24Bdi8Gd48bSPMeeGW48c8ZMsnebRO7G22kGQC3JxmYHcGiRSGOQnMF2mHHjjBy2G7wgLbMTQFoOJG7DoUV+Rhrzzxn/DttvnJH+7ObfbRLphrPTP+DVwnAjjU3iY9vhxPkSCWa3gbYkyEvn4LfFAKjF4mPf4eQNPGfMf8tukzDcIJ1TcCDBALdfQA67kfDtsO389vbHhm+31cnLz07f/OFDhZ0cLi0I6w6gMAwIKAdb14DOGAWjYBSMglEABQBUlmXB2BC0LgAAAABJRU5ErkJggg==","orcid":"","institution":"Wildlife Institute of India","correspondingAuthor":true,"prefix":"","firstName":"Shravana","middleName":"","lastName":"Goswami","suffix":""},{"id":400510524,"identity":"93748c64-d628-4a3e-8ee7-3783d2b83a7e","order_by":1,"name":"Ujjwal Kumar","email":"","orcid":"","institution":"Wildlife Institute of India","correspondingAuthor":false,"prefix":"","firstName":"Ujjwal","middleName":"","lastName":"Kumar","suffix":""},{"id":400510525,"identity":"a9853782-0023-4e3d-9e81-eeec728c8ec0","order_by":2,"name":"Qamar Qureshi","email":"","orcid":"","institution":"Wildlife Institute of India","correspondingAuthor":false,"prefix":"","firstName":"Qamar","middleName":"","lastName":"Qureshi","suffix":""},{"id":400510527,"identity":"fe118889-6f09-46b3-99a3-dfc4d6120186","order_by":3,"name":"Yadvendradev Jhala","email":"","orcid":"","institution":"Wildlife Institute of India","correspondingAuthor":false,"prefix":"","firstName":"Yadvendradev","middleName":"","lastName":"Jhala","suffix":""}],"badges":[],"createdAt":"2025-01-03 07:08:07","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5756049/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5756049/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-08093-0","type":"published","date":"2025-07-02T15:57:56+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":73686090,"identity":"1745ddb3-e531-4ca3-93bb-f81640c8698d","added_by":"auto","created_at":"2025-01-13 14:40:15","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":235866,"visible":true,"origin":"","legend":"\u003cp\u003eSeasonality of breeding and fawning in chital (Axis axis) as indexed by proportion of different antler stages of males and lactating females in relation to resource availability indexed by NDVI in Kanha Tiger Reserve.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-5756049/v1/159134ec372575b305a0bbc0.png"},{"id":73686083,"identity":"06e6f04c-0b43-4ffa-8ef9-6200d7cb092e","added_by":"auto","created_at":"2025-01-13 14:40:14","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":28316,"visible":true,"origin":"","legend":"\u003cp\u003ePercent tiger litter weaned (n=31) depending on their birth month data obtained from Kumar, 2019 vs proportion of lactating chital females in chital population in Kanha tiger reserve every month. Blue bars show the number of tiger litter and orange line show the percentage of lactating chital females in population.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-5756049/v1/1b4e73d312dcc429004f64fa.png"},{"id":86179141,"identity":"8af4d9a4-2948-4f0a-b6d7-4b91852c3236","added_by":"auto","created_at":"2025-07-07 16:16:17","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1725467,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5756049/v1/466a01f8-8b40-490b-bd0c-728a2aae7200.pdf"},{"id":73686080,"identity":"cf0b1070-b915-4e42-bcbd-4855e2b4e504","added_by":"auto","created_at":"2025-01-13 14:40:13","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":688114,"visible":true,"origin":"","legend":"","description":"","filename":"Suppliment.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5756049/v1/c16c04efac4c0dc222a69673.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Ecological Determinants of Chital (Axis axis) Reproductive Seasonality in Kanha Tiger Reserve","fulltext":[{"header":"Introduction","content":"\u003cp\u003eSynchrony or birthing peaks are seen as a response to either seasonal environments or as an anti-predatory adaptation (Roeder et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Tropical and equatorial mammals exhibit a lack of seasonality in breeding due to uniform resource availability. However, tropical India is highly seasonal in resource availability due to the rainfall being primarily restricted to the monsoon season. Sinclair et al. (\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) studied 13 African ungulates and found that their breeding synchronizes with seasons of abundant forage resources, an adaptation that enhances offspring survival by ensuring that young are born during favourable conditions. Rainfall patterns also significantly influence ungulate reproductive behaviour; Priyadarshini (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) notes that rainfall affects food availability, which in turn determines reproductive success. Increased rainfall can lead to lush vegetation, providing better nutrition for pregnant or lactating females and improving offspring survival rates. The reproductive patterns of ungulates can vary significantly based on environmental conditions. Ungulates are timed to coincide with their breeding season with periods of highest plant quality to support offspring nutrition (Mduma et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). This adaptation ensures that lactating females and weaned offspring obtain the required energy and protein from the forage (Western \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Bronson \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Owen-Smith et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Priyadarshini \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). These relationships underscore the importance of understanding ecological dynamics to better manage ungulate populations and their habitats. As environmental conditions continue to change due to human impact, ongoing research will be required to understand how ungulates adapt to these challenges. Changes in forage quantity and quality significantly impact the body condition of ungulates, especially females during gestation and lactation, when their energy and protein requirements are heightened. Studies by Oftedal \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Robbins \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Parker et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Gaillard et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Sinclair et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; and Cook et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2001\u003c/span\u003e all emphasize the critical role that adequate nutrition plays in ensuring the health and reproductive success of these animals during these demanding periods. The onset of oestrus in females is significantly affected by conditions required for a successful pregnancy and subsequent lactation, such as food availability or favourable weather conditions (McGinnes \u0026amp; Downing \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1977\u003c/span\u003e; Asher \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), as increased time and energy expenditures for food searching can lead to higher mortality in reproducing adults (Robbins \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e1993\u003c/span\u003e). We hypothesize that after a stressful dry summer, the new flush in vegetation caused by the monsoon rains should be the cue for ovulation in female ungulates, including chital in India.\u003c/p\u003e \u003cp\u003eIn most of the cervids, antler cycles serve as external indicators of internal reproductive changes (Bubenik et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). In most deer species, males undergo a yearly cycle characterized by the growth and subsequent shedding of antlers (Bubenik et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Tom\u0026aacute;s \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Ungerfeld et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Antlers play a crucial role in male-male confrontations to establish dominance and secure mates, serving as an indication to females of male fitness and condition during the rutting season, which is a period of intense mating activity (Bubenik et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Clements et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Vanp\u0026eacute; et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Heckeberg \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Testosterone levels peak seasonally during the rut season (Bubenik \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1990\u003c/span\u003e), and when testosterone levels fall below threshold after the rutting season, antlers are cast off in most cervids (Putman \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Bartoš \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Bubenik \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). Consequently, peak rutting activity often coincides with the period when the largest proportion of males exhibit burnished antlers, indicative of their reproductive readiness (Raman \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Ramesh et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Bhusal et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In deer, observing actual fawning is a rare event and is often studied by implanting radio transmitters in the vagina of pregnant females, which are expelled out at the time of parturition (Bowman \u0026amp; Jacobson \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Conant et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This allows for the study of timing and specific location of parturition. We indexed the timing of fawning by estimating the proportion of lactating females in the population (Priyadarshini \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Jhala \u0026amp; Isvaran \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMany studies on breeding seasonality have been conducted on North American and African ungulates, but in the Indian subcontinent only a few ungulate species have been studied for their reproductive patterns (Rice \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Raman \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). However, these studies have primarily focused on observing the seasonality of the antler cycle. Management decisions like prey augmentation (Phumanee et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), where animals from high-density areas were captured and moved to low-density areas to increase the carrying capacity of large carnivores, also need information about rutting and fawning. Translocation during prey augmentation in the fawning season can cause stress due to possible mother-fawn separation, and aggressive behaviour of males that are in hard antlers during the rutting season can increase mortality, making it essential to understand the breeding season of the source chital population beforehand. In this study, we record the reproductive condition of chital throughout the year from representative samples to decipher seasonality and its likely drivers in the dry deciduous forest system of Kanha National Park.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eKanha Tiger Reserve is located in the Satpura Maikal hill ranges of the Central Indian highlands (Rodgers \u0026amp; Panwar \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1988\u003c/span\u003e). It was among the first nine Tiger Reserves gazetted in 1973. It is located between 80\u0026deg;26\u0026rsquo;10\u0026rsquo;\u0026rsquo; E to 81\u0026deg;04\u0026rsquo;40.00\u0026rsquo;\u0026rsquo;E and 22\u0026deg;01\u0026rsquo;5.0\u0026rdquo; N to 27\u0026deg;27\u0026rsquo;48.00\u0026rdquo; N in Balghat and Mandla districts of Madhya Pradesh. The area of the Core Zone is 917.43 km2, and that of the Buffer Zone is 1134.39 km\u003csup\u003e2\u003c/sup\u003e, adding up to 2051.82 km\u003csup\u003e2\u003c/sup\u003e as the total area of Kanha Tiger Reserve. The Tiger Reserve has flat hilltops, varying degrees of slopes, and meadows in the valleys, which offer unique, diverse types of wildlife habitat, forming ideal niches for various species of plants and animals.\u003c/p\u003e \u003cp\u003eThere are three distinct seasons in Kanha Tiger Reserve. These are monsoon, summer, and winter. Summer extends from mid-February to June end, monsoon is from June end week to mid-October, and winter is from mid-October to mid-February. Due to a great variation in temperature, humidity, wind velocity, and precipitation in different seasons, these factors serve as regulators of vegetation and the habits of wild animals in the reserve. Kanha Tiger Reserve lies in a dry deciduous zone, and the availability of water is a very important factor for the survival of animals and plants. Major forest types of the area can be divided into four categories: sal forest, miscellaneous forest, bamboo mixed forest, and grasslands. Controlled burning practices during February are carried out in the grasslands of Kanha each year for habitat improvement and management purposes.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eField methods\u003c/h3\u003e\n\u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eEstimating proportion of population in breeding condition\u003c/h2\u003e \u003cp\u003eObservation of actual mating events and fawning in wild populations is rare, especially in tropical forest systems. We therefore indexed breeding condition in males by assessing their antler condition (Bubenik et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ramesh et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and fawning in females by assessing their lactation state (Fryxell \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Eberhardt et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Laurian et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Priyadarshini \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor obtaining an unbiased representation of the population in samples, we conducted intensive surveys along fixed vehicle routes to observe the breeding status of male and female chital groups. Each route was visited once a week. Initially, the population was categorized into adults, sub-adults, and juveniles, following Schaller (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e1967\u003c/span\u003e). For assessing breeding signs, only adult individuals were considered. To assess the involvement of sub-adult females in breeding, a total of 200 sub-adult females were observed for lactation signs during the peak lactation period. We subdivided adult males into hard antler, velvet, and shed antler male groups as per their antler stages at different times of the year (Figure \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). We expect that the peak breeding season will coincide with the time when the largest proportion of bucks exhibit hard antlers.\u003c/p\u003e \u003cp\u003eTo determine the fawning season, lactation signs were recorded whenever an adult female was observed, as the presence of lactating mothers indicated the birth of a fawn (Fryxell \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Eberhardt et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Laurian et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Priyadarshini \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). In chital udders and teats become visible only a few days before parturition and remain prominent up to four months after parturition (Weigl \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Myhrvold et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) (Figure S2).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eAnalytical method\u003c/h3\u003e\n\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eMating and fawning season\u003c/h2\u003e \u003cp\u003eData on various stages of male antlers and lactating females were pooled month-wise for exploratory analysis. Then the data were categorized into three distinct seasons: winter, summer, and monsoon, and reanalysed. Since the same route was followed weekly for observations, the computation was done using multiple sampling with replacement approach (Skalski et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eNDVI change\u003c/h2\u003e \u003cp\u003eChital are the primary grazers at Kanha Tiger Reserve as more than 50% of their diet consists of grasses (Awasthi \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). So change in the condition of grass affects the body requirements of chital. To see the monthly productivity in the grasslands of Kanha, we analysed the Normalized Difference Vegetation Index (NDVI) of the grasslands of Kanha for the past 10 years from satellite imagery of LANDSAT 8 (USGS) in Google Earth Engine (Gorelick et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). A harmonic model for monthly change in NDVI was fitted to the data to see monthly changes in the grasslands of Kanha (Fig S3). When sprouting begins in grasslands, the NDVI value starts increasing from a low as the grasses turn green replacing the dry grass or bare land left after burning. Then monthly average values of NDVI were plotted to evaluate the synchrony between grass sprouting season and chital breeding in Kanha.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eBirth of tiger cubs\u003c/h3\u003e\n\u003cp\u003eWe used information from Kumar (\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) on the number of litters born each month for tigers in Kanha. Kumar (\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) determined the birth month of tiger litters by aging the cubs. The birth records and the month of parturition were obtained through continuous sightings by experienced forest staff, researchers, camera trap pictures, and handheld photographs taken by tourists who shared the date and location information. Information on 31 litter births was recorded with reasonable certainty. Once the month of birth was ascertained, the relationship between chital fawning and tiger cub was explored by looking at the time of birth of offspring of both prey and predator.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eFawning occurred throughout the year as lactation was recorded in all months. However, the proportion of lactating females in the population peaked in March-April. April has the highest proportion of lactating females (78\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8%), followed by March (73.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1%), while November recorded the lowest proportion (6.93\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37%) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Seasonally, summer had the highest proportion of lactating females (72.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24%), followed by monsoon (48.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46%), and winter (30.79\u0026thinsp;\u0026plusmn;\u0026thinsp;0.34%) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). None of the 200 sampled sub-adult females were observed to be lactating.\u003c/p\u003e \u003cp\u003eIn July, nearly all adult males have hard antlers (97.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15%). Between May and November, over 60% of the male population displayed hard antlers, while more than 50% of the adult male population sprouted velvet-covered antlers between December and April (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, table S2). The proportion of shed antler male observation is low due to the short interval between antler shedding and the emergence of new antlers.\u003c/p\u003e \u003cp\u003eIn Kanha Tiger Reserve, grassland growth peaks during the monsoon and post-monsoon periods but declines during the dry winter season with minimal rainfall. The NDVI value during the post-fire season (April) shows an upward trend indicating the emergence of freshly grown grass shoots. Lactating peaked at the time of the new grass flush when the NDVI values start having an upward trend (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table S3) after a lowest in March. Births of tiger litters were observed throughout the year with a peak in December. Their weaning started after 3\u0026ndash;4 months which coincided with the same time of fawning (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThere seems to be a debate regarding the existence of distinct seasonality in chital, particularly in tropical regions where food and water are relatively stable throughout the year. Krishnan (\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1972\u003c/span\u003e) states that chital lacks any birth peak throughout peninsular India, but many studies found distinct seasonal peaks in chital rutting behaviour in India (Table S4). Studies were done in Hawaii by Graff \u0026amp; Nichols (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1966\u003c/span\u003e), Texas by Ables (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1977\u003c/span\u003e), Australia by Kelly et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), Nepal by Bhusal et al. (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and Mishra (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1982\u003c/span\u003e) also found a distinct peak in rutting behaviour of chital depending on environmental factors like day length and resource availability. The Central Indian landscape has distinct seasonality, which affects the availability and quality of forage during different times of the year. Mixed foragers like Chital shift their diet towards browsing during summer when grass nutrient quality declines (Awasthi \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Watter et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), emphasizing the importance of dietary quality. The dry grasses become non-nutritious and are filled with silica and lignin content (Kayongo-Male et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1976\u003c/span\u003e). Due to the burning of grasslands during February to reduce the fuel load of grasses, preventing uncontrolled fire during the summer season, fresh biomass appears post-fire season, which is high in crude protein (White \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Akbarinia \u0026amp; Koocheki \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Arzani et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1998\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and other minerals like calcium and magnesium (Devi et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). We found that the breeding peak of chital is synchronous with the appearance of fresh grass forage biomass, i.e., the months of March and April. Moreover, fawns born at this time were weaned off during the monsoon period, when there is ample forage available for them to feed on. Ogutu et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) noticed the birth peak of ungulates coincided with seasonal rainfall and the appearance of green foliage in tropical and subtropical regions. Clutton-Brock et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1989\u003c/span\u003e) also state that ungulate breeding is affected by food availability, and this hypothesis is also supported by Ahrestani et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), who conclude that chital also follows this pattern.\u003c/p\u003e \u003cp\u003eIn Kanha, rutting occurs in late summer and monsoon season, although some hard antler males are present in the population throughout the year. Deer females conceive only when they have good body condition (Flajšman et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Paoli et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), and Kelly et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) found that this is true for chitals also. In Kanha, chital has better body condition when ample food is available (Awasthi \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Similarly, fawning starts in January and reaches a peak during April. Since fawning is represented by lactating females and lactation occurs up to 4 months in chitals, it is difficult to determine the exact fawning peak.\u003c/p\u003e \u003cp\u003ePeak in tiger cub birth is the same time when more than half of the female chital were lactating. When weaning of tiger cubs starts after a few months, it is the same time when the number of lactating females is also highest. When weaning starts, the need for prey also increases, as the mother needs to provide food for weaned cubs. During this period, chitals are most vulnerable to predation, with males being in hard antlers and preoccupied with the rut, females caring for fawns and weak due to lactation, and ample availability of fawns. Tigers in the Kanha landscape showed a functional response by selective predation on the abundant chital fawns and rutting males (Jhala et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), resulting in numerical response by timing their birth peak at this time. Similar pattern has also been reported in mountain lions (\u003cem\u003ePuma concolor\u003c/em\u003e) by Jansen \u0026amp; Jenks (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). The mountain lion birth peak is synchronized with the birth of North American ungulates in May-June.\u003c/p\u003e \u003cp\u003eThere are certain cues like resource abundance that track ecological rhythms that influence reproductive behaviour. The intricate relationship between seasonal forage dynamics, reproductive patterns, nutritional ecology, and predation underscores the importance of understanding these dynamics for effective wildlife management and conservation planning. Birthing and rutting peaks of chital in Kanha, which is a deciduous forest-grassland system, were determined by resource availability and were unlikely to be in response to predation.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study on the seasonality of chital, particularly in tropical regions with stable food and water availability, highlights the complexity of their reproductive behaviour.\u003c/p\u003e \u003cp\u003eIn the central Indian landscape, seasonality significantly affects forage availability and quality. The management practices in tiger reserves, such as controlled burning, enhance forage quality by promoting the growth of nutrient-rich fresh biomass. This study finds that the breeding peak of chital in Kanha synchronizes with the appearance of fresh grass forage in March and April after controlled burning, providing crucial nutrition to lactating females. Rutting occurs in late summer and monsoon, while fawning peaks in April, which represents a proxy for the presence of lactating females, making them and their fawns more vulnerable to predation. The alignment of tiger reproductive biology with the peak availability of chital fawns suggests a functional predator response.\u003c/p\u003e \u003cp\u003eThe interplay between seasonal forage dynamics, reproductive patterns, nutritional ecology, and predation is vital for effective wildlife management and conservation. Understanding these dynamics in the deciduous forest-grassland system of Kanha reveals that chital birthing and rutting peaks are driven more by resource availability than predation pressures. Effective conservation strategies must consider these intricate ecological relationships to support the sustainable management of chital populations and their habitats.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Chief Wildlife Warden of Madhya Pradesh and management of Kanha Tiger Reserve for permissions and logistics for the study. We thank our field assistants Nirottam and Kanhaiya and the team of forest guards for their help in field data collection. We thank field director Kanha, S.K. Singh for logistic support\u003cstrong\u003e.\u0026nbsp;\u003c/strong\u003eThis study was funded by National Tiger Conservation Authority, Ministry of Environment, Forest and Climate change, Government of India.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors' contributions:\u003c/strong\u003e YVJ \u0026amp; QQ conceived, supervised, procured the resources for the study, SG did the field data collection, SG \u0026amp; YVJ did the data analysis, SG, UK and YVJ wrote the MS, and all the authors approved the MS.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and material:\u0026nbsp;\u003c/strong\u003eAll data generated and analysed during this study are included in this published article and its supplementary information files.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval:\u0026nbsp;\u003c/strong\u003eThe study was based on observation. All observation was made in the animal’s natural habitat without any physical interference. The study complied with the ethical guidelines issued by Chief Wildlife Warden through permit No. Serial no./D.M.II/Research-213/9027.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests:\u0026nbsp;\u003c/strong\u003eThe authors declare that they have no conflict of interest.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003e\u003cstrong\u003eAbles, E.D. (1977). \u003c/strong\u003e\u003cem\u003eThe axis deer in Texas \u003c/em\u003eVol. 2. College Station: Texas A\u0026amp;M University Press.\u003c/li\u003e\n\u003cli\u003e\u003cstrong\u003eAhrestani, F.S., Van Langevelde, F., Heitk\u0026ouml;nig, I.M.A., \u0026amp; Prins, H.H.T. 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(2020). \u003c/strong\u003eSeasonal diet preferences of chital deer in the northern Queensland dry tropics, Australia. \u003cem\u003eThe Rangeland Journal\u003c/em\u003e 42(3): 211.; https://doi.org/10.1071/RJ20015\u003c/li\u003e\n\u003cli\u003e\u003cstrong\u003eWeigl, R. (2005). \u003c/strong\u003e\u003cem\u003eLongevity of mammals in captivity: from the living collections of the world ; a list of mammalian longevity in captivity\u003c/em\u003e. Schweizerbart, Stuttgart, 214pp.\u003c/li\u003e\n\u003cli\u003e\u003cstrong\u003eWestern, D. (1979). \u003c/strong\u003eSize, life history and ecology in mammals. \u003cem\u003eAfrican Journal of Ecology\u003c/em\u003e 17(4): 185\u0026ndash;204.\u003c/li\u003e\n\u003cli\u003e\u003cstrong\u003eWhite, L.M. (1983). \u003c/strong\u003eSeasonal changes in yield, digestibility, and crude protein of vegetative and floral tillers of two grasses. \u003cem\u003eJournal of Range Management\u003c/em\u003e 36: 402\u0026ndash;404.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Fawning, Lactation, Rutting, Nutrition, Tiger Births","lastPublishedDoi":"10.21203/rs.3.rs-5756049/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5756049/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eReproductive seasonality in mammals is often influenced by environmental factors like resource availability, predation. While tropical and equatorial regions exhibit continual breeding due to stable resources, tropical India experiences pronounced seasonality in forage availability due to monsoonal rainfall. This study investigates the breeding timing of chital (\u003cem\u003eAxis axis\u003c/em\u003e) in the tropical deciduous forests of Central India at Kanha Tiger Reserve, where forage availability varies seasonally. Field observations focused on antler conditions in males and lactation signs in females to estimate breeding timing. Rutting peak, as determined by males with hard antlers, coincided with nutritional peak induced by monsoon, which probably was the cue for ovulation in females. Lactation, the most nutritionally demanding period, peaked with the post-fire sprouting of herbaceous vegetation in April and fawn weaning synchronized with ample food availability. This synchrony between reproductive cycles and forage availability emphasizes the adaptive strategies of chital to maximize offspring survival. Furthermore, the peak in tiger births (54%) coincided with the fawning peak of chital. making them vulnerable prey during a nutritionally demanding period for tigresses, who are restricted in their movements near vulnerable cubs. Understanding the links between trophic levels and their seasonality offers important insights for wildlife management and conservation strategies.\u003c/p\u003e","manuscriptTitle":"Ecological Determinants of Chital (Axis axis) Reproductive Seasonality in Kanha Tiger Reserve","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-01-13 14:39:47","doi":"10.21203/rs.3.rs-5756049/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-04-29T04:17:25+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-24T06:26:13+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-16T06:36:06+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-15T10:06:03+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-14T18:10:27+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"193286081361586534898174084403127698337","date":"2025-04-07T10:49:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"206611647173081804409717130598192835223","date":"2025-04-04T12:38:07+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"301723618395966852739102537778967656308","date":"2025-04-02T08:38:28+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"240354654899649674398921079352620121237","date":"2025-04-02T07:43:35+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-04-02T07:31:22+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-02-25T08:24:52+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-01-13T15:46:41+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-01-10T13:48:21+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-01-03T06:52:38+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"b28256a1-b646-4572-959d-5234162e7dc8","owner":[],"postedDate":"January 13th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":42688241,"name":"Biological sciences/Ecology"},{"id":42688242,"name":"Biological sciences/Zoology"},{"id":42688243,"name":"Earth and environmental sciences/Ecology"},{"id":42688244,"name":"Earth and environmental sciences/Environmental sciences"}],"tags":[],"updatedAt":"2025-07-07T16:04:33+00:00","versionOfRecord":{"articleIdentity":"rs-5756049","link":"https://doi.org/10.1038/s41598-025-08093-0","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2025-07-02 15:57:56","publishedOnDateReadable":"July 2nd, 2025"},"versionCreatedAt":"2025-01-13 14:39:47","video":"","vorDoi":"10.1038/s41598-025-08093-0","vorDoiUrl":"https://doi.org/10.1038/s41598-025-08093-0","workflowStages":[]},"version":"v1","identity":"rs-5756049","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5756049","identity":"rs-5756049","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}