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Divided They Stay : Species Coexistence In A Community Of Mutualists And Exploiters

2012-12-14T07:02:51Z

Title: Divided They Stay : Species Coexistence In A Community Of Mutualists And Exploiters Authors: Ghara, Mahua Abstract: The fig–fig wasp interaction is a classic example of obligate mutualism and coevolution. It is also a nursery pollination mutualism and supports a diversity of exploiter/parasite/non-pollinating fig wasp species. Mutualists and exploiters comprise the fig wasp community. All the wasp community members are obligately dependent on the fig syconium (a globular closed structure comprising of hundreds to thousands of uniovulate florets) for completing their life cycle. The fig florets can be sessile (without a stalk) or pedicellate (stalked) and can support a community comprising 3–30 wasp species. Fig wasps can access the floral resources for oviposition directly by entering into the syconium (internal oviposition) or by penetration of the syconium surface (external oviposition). Most studies on the fig–fig wasp interaction have investigated the stability of the interaction, pollination biology, pollen dispersal, co-evolution or the effect of exploiters on this mutualism. However, studies dealing with community ecology and species coexistence mechanisms in these communities are rare. Factors contributing to coexistence of mutualists and exploiters in a fig wasp community were studied using a reasonably speciose fig wasp community associated with Ficus racemosa in south India. The wasp community of Ficus racemosa comprises a single species of pollinator and six species of exploiters; together they represent three genera of fig wasp species. The community members show differences in their feeding habit; they could be 1) gallers (feed on floral tissue after pollination and/or after inducing abnormal tissue development of the floret that is also called the gall), 2) inquilines (feed on gall tissue but cannot induce galling; survive by feeding on gall tissue and starving the host larva to death), or 3) parasitoids (lay eggs in or on developing offspring of a galler or inquiline species; develop by feeding on host tissue). Resource partitioning across temporal and spatial axes on this fig wasp community have been quantified. Ovipositor traits of each community member were also investigated since variation in ovipositor traits might facilitate resource partitioning. Finally, the role of life-history traits in species coexistence in this community was also explored. Temporal resource partitioning among members of the fig wasp community was studied (1) across the resource phenology, i.e. over the development phases of the fig syconium, and (2) on a diel scale. The seven members of the wasp community were found to partition their oviposition periods across fig syconium development phenology; some species used very young syconia (soft and smaller in size) for oviposition whereas others used mature (hard and bigger in size) syconia for oviposition. The first species to colonise the syconia were gallers and these were followed by parasitoids in a definite oviposition sequence. Pollinators arrived concurrently with an exploiter galler species and had the shortest oviposition window in terms of days. Although fig wasps are known to be largely diurnal, night oviposition in several fig wasp species was documented for the first time. Wasp species showed a peak in their activity period across the diel cycle and phenology. This is probably the first study to simultaneously investigate temporal partitioning across the syconium phenology as well as the diel scale in a fig wasp community. Partitioning of syconium space was investigated by quantifying the quality (type of floret—sessile or pedicellate) of floral resources. The number of individuals of each species developing in a syconium was quantified along with host accessibility during oviposition by each wasp species. The association between community members developing within a syconium was also tested. The differential occupancy of florets by each species based on their distance from the base of the syconium was evaluated. For the first time the relative distribution of males and females of the entire fig wasp community was quantified. The wasp community members used similar types of florets for oviposition. Seeds were found mostly in sessile florets and wasps were present in large numbers in pedicellate florets. Except for one wasp species, all others occurred uniformly within the syconium with respect to the distance from the base of the syconium. Species distribution models revealed higher prediction ability for the location of mutualists (seed and pollinator) within the syconium compared to exploiters. Within a syconium, all species pairs exhibited positive associations indicating either an absence of or low competitive exclusion. Some florets were modified by their gall occupants such that they were longer in length indicating the possibility of creation of an enemy-free zone by the gall occupant. Yet, most florets were accessible to ovipositing wasps based on ovipositor lengths and flexibility. The probability of finding a male decreased with increase in floret length when all wasp species were grouped together; however, this trend did not hold true when males and females of species were tested individually. Based on these results, the fig wasps of F. racemosa could be grouped into—(1) Early-arriving galler species which used immature florets, inducing large galls that protruded into the cavity, and with fewer individuals per syconium, (2) Galler species arriving concurrently with the pollinator, inducing galls that were morphologically indistinguishable from those of the pollinator, and with many individuals developing per fig syconium, and (3) Parasitoids and/or inquilines of the galler species, with variable abundance per syconium. Thus, these results show that the wasp species do not clearly partition floral resources between syconia and within syconium but they can modify their oviposition sites and also differ in the proportion of florets within a syconium used for oviposition. Oviposition sites of the fig wasps can be reached only by using their ovipositor. The resources for oviposition are hidden and hence might require tools for resource location and utilisation. The frequency and diversity of sensilla on the ovipositor, as well as ovipositor structure (morphology and sclerotisation of the tip) was documented for the entire wasp community. The internally-ovipositing pollinator had the simplest ovipositor, negligible sclerotisation and only one type of sensillum on its ovipositor; the externally ovipositing exploiter species had teeth on their ovipositors, sclerotisation and various types of ovipositor sensilla. Ovipositor sclerotisation and lateness of arrival for oviposition in syconium development were positively correlated. Ovipositor teeth height increased from gallers to parasitoids. Presence of different types of sensilla was noted which included mechano- and chemosensilla, as well as combined mechano-chemosensilla. Chemosensilla were most concentrated at ovipositor tips while mechanoreceptors were more widely distributed. Ovipositor traits of one putative parasitoid/inquiline species differed from those of its syntopic galler congeners and clustered with those of parasitoids within a different wasp subfamily. Thus ovipositor tools show lability based on adaptive necessity, and are not constrained by phylogeny. Life-history traits such as fecundity, pre-adult and adult lifespan were studied for each wasp member of the community. Trade-offs in life-history traits were also investigated. Interspecific variation in life-history traits was observed. Gallers were pro-ovigenic (all eggs were mature upon adult emergence) whereas parasitoids were synovigenic (eggs matured progressively during adult lifespan). Initial egg load was correlated with body size for some species, and there was a trade-off between egg number and egg size across all species. Although all species completed their development and left the syconium concurrently, they differed in their adult and preadult lifespans. Providing sucrose solutions increased parasitoid lifespan but had no effect on the longevity of some galler species. While feeding regimes and body size affected longevity in most species, an interaction effect between these variables was detected for only one species. Life-history traits of wasp species exhibited a continuum in relation to their arrival sequence at syconia for oviposition during syconium development, and therefore reflected their ecology. The largest number of eggs, smallest egg sizes, and shortest longevities were characteristic of the earliest-arriving galling wasps at the smallest, immature syconia; the converse characterised the later-arriving parasitoids at the larger, already exploited syconia. Thus life-history is an important correlate of community resource partitioning and can be used to understand community structure. The comparative approach revealed constraints and flexibility in trait evolution. This is probably the first comprehensive study of life-history traits in a fig wasp community.

Phylogeny And Biogeography Of Hemidactylus Oken, 1817 Geckos Of The Indian Subcontinent

2013-02-25T09:38:19Z

Title: Phylogeny And Biogeography Of Hemidactylus Oken, 1817 Geckos Of The Indian Subcontinent Authors: Bansal, Rohini Abstract: Geckos of the genus Hemidactylus Oken, 1817 are one of the most widely distributed and species-rich groups of gekkonids. With approximately 90 species, the genus Hemidactylus accounts for 10% of the total number of species in the family Gekkonidae (Carranza and Arnold, 2006). They are found in a range of ecological conditions from wet tropical forest to arid deserts. Their wide distribution is believed to have been facilitated not only naturally, but also through human mediated transport. Indian subcontinent houses 31 species of these geckos, 23 of which are endemic to the subcontinent. The aim of this study was to understand the phylogeny and biogeography of Hemidactylus geckos of the Indian subcontinent. In order to arrive at the conclusions regarding the biogeographic history of this widely distributed taxon on the Indian subcontinent, an integrative approach was followed. First the phylogenetic reconstruction of Indian species was undertaken. The ancestral areas of distribution were reconstructed on the phylogeny and the divergence dates of the species were also estimated. Finally, the biogeographic events were inferred in accordance with the geological events with respect to the divergent times for these radiations. Results indicated that Hemidactylus geckos of the Indian subcontinent belong to three geographical clades: Southeast Asian clade, West-Asian arid clade and a ‘unique Indian radiation’ which consisted majority of the species distributed in India that are largely confined to the Indian subcontinent. Additionally, the three widely distributed, commensal species (H. brookii, H. frenatus and H. flaviviridis) were nested within the Indian radiation suggesting their Indian origin. It was also seen that the endemic Hemidactylus geckos of Sri Lanka have been derived from India through multiple independent dispersal events spread across the Oligocene-Miocene. These dispersals were both geo dispersal as well as transmarine dispersal events. However, the species distributed both in India and Sri Lanka appear to have undergone recent dispersals into Sri Lanka and probably back into India, primarily through human agency. Thus, it was inferred that the Indian subcontinent has served as an important arena for diversification among the Hemidactylus geckos and their spread. In addition, it was discovered that H. anamallensis, a species endemic to Southern Western Ghats of peninsular India was probably genetically distinct from Hemidactylus, indicating that it could not be Hemidactylus at all. Thus, the phylogenetic position of H. anamallensis within the subfamily Gekkoninae was also investigated and its genetic distinctiveness from that of Hemidactylus was also tested. Results showed that H. anamallensis lineage was indeed distinct from Hemidactylus group, as well as from other closely related genera (Cyrtodactylus and Geckoella) in both nuclear and mitochondrial markers. Divergence estimates supported a scenario wherein H. anamallensis dispersed across marine barrier to the drifting peninsular Indian plate in the late Cretaceous, whereas Hemidactylus arrived on the peninsular India after the Indian plate collided with the Eurasian plate. Based on these molecular evidence and biogeographical scenario it was proposed that genus Dravidogecko should be resurrected.

Ants, Figs, Fig Wasps : The Chemical Ecology Of A Multitrophic System

2013-04-01T07:20:53Z

Title: Ants, Figs, Fig Wasps : The Chemical Ecology Of A Multitrophic System Authors: Ranganathan, Yuvaraj Abstract: Plant–animal interaction systems are complex food webs where the members—plants, pollinators, herbivores, parasites and predators of the pollinators/herbivores—interact with each other in ways which maximize their own fitness. Based on the net outcome, such interactions could be mutually beneficial to the interacting members (mutualism) or beneficial to only one of the interacting members at the cost of the other interacting members (herbivory, predation, parasitism). It is possible that such outcomes are actually a continuum and could swing in either direction from beneficial to detrimental and vice versa. Such transitions happen not only over long time scales, but could also happen within shorter time scales based on conditionalities. Conditional outcomes are those in which the outcome of an interaction between two partners is conditional on the involvement of a third partner. Thus, studying such outcomes necessitates taking into account systems beyond the classical two-partner interactions. In such complex multitrophic plant–animal interaction systems in which there are direct and indirect interactions between species, comprehending the dynamics of these multiple partners is very important for an understanding of how the system is structured. In Chapter 2 we investigate Ficus racemosa and its community of obligatory mutualistic and parasitic ﬁg wasps that develop within the fig inﬂorescence or syconium, as well as their interaction with opportunistic ants. We focus on temporal resource partitioning among members of the ﬁg wasp community over the development cycle of the fig syconia during which wasp oviposition and development occur and we study the activity rhythm of the ants associated with this community. We found that the members of the wasp community partitioned their oviposition across fig syconium development phenology and showed interspecific variation in activity across the diel cycle. The wasps presented a distinct sequence in their arrival at fig syconia for oviposition. We documented night oviposition in several fig wasp species for the first time. Ant activity on the fig syconia was correlated with wasp activity and was dependent on whether the ants were predatory or trophobiont-tending species; only numbers of predatory ants increased during peak arrivals of the wasps. In Chapter 3, we found that predatory ants (Oecophylla smaragdina) patrolling F. racemosa trees were attracted to the odour from fig syconia at different developmental phases, as well as to the odours of fig wasps, whereas other predatory ants (Technomyrmex albipes) responded only to odours of syconia from which ﬁg wasps were dispersing and to ﬁg wasp odour. However, trophobiont-tending ants (Myrmicaria brunnea) patrolling the same trees and exposed to the same volatiles were unresponsive to ﬁg or ﬁg wasp odours. The predatory ants demonstrated a concentration-dependent response towards volatiles from figs receptive to pollinators and those from which wasps were dispersing while the trophobiont-tending ants were unresponsive to such odours at all concentrations. Naıve predatory ants failed to respond to the volatiles to which the experienced predatory ants responded, indicating that the response to ﬁg-related odours is learned. In Chapter 4 we characterise the dynamics of the volatile bouquet of the ﬁg syconium from the initiation through pre-receptive, receptive, and late inter-ﬂoral stages which act as signals/ cues for different ﬁg wasp species. We were also interested in diel patterns of volatile emission as some ﬁg wasp species were strictly diurnal (the pollinator, Ceratosolen fusciceps) whereas other ﬁg wasps such as Apocryptophagus fusca were observed ovipositing even during the nocturnal hours. We identified volatiles that were specific to syconium development phase as well as to the time of day in this bouquet. α-muurolene was identified as the sesquiterpene specific to receptive-phase as well as being present only during the day thus coinciding with the diurnal pollinator arrival pattern. Volatiles such as (E)-β-ocimene were present in increasing levels across the developmental stages of the ﬁg and thus could act as background volatiles providing suitable information to fig wasps about host plants and their phases. Chapter 5 examines the responses of predatory and trophobiont-tending ant species to the cuticular hydrocarbon (CHC) extracts of four galler and two parasitoid ﬁg wasp species associated with F. racemosa. Interestingly, the antennation response of both experienced and na¨ıve ants to these wasp extracts was identical indicating that prior exposure to such compounds is not necessary for eliciting such response. We also characterised these cuticular hydrocarbon extracts to find potential compounds which could as short-range cues for predatory ants. Ants were more responsive to the cuticular extracts of parasitoids rather than to those of galler wasps, implying that the CHC profile of carnivorous prey may contain more elicitors of aggressive behaviour in ants compared to herbivorous prey whose profiles may be more similar to those of their plant resources. We also find congruency between the cuticular profiles of parasitoids and their hosts suggesting that parasitoids could sequester compounds from their diet. Important findings and conclusions of the thesis are presented in Chapter 6. The first two parts of the appendices section discuss work carried out on alternative ways of analysing multivariate data sets such as plant volatiles and insect cuticular hydrocarbons. Appendix A details the use of Random Forests, an algorithm-based method of analysing complex data sets where there are more variables than samples, a situation akin to microarray data sets. This work illustrates the use of such techniques in chemical ecology, highlighting the potential pitfalls of classical multivariate tests and the advantages of newer more robust methods. Appendix B, an invited article following the publication of the earlier work, compares different data transformation procedures currently employed in such multivariate analysis. Appendix C details sex-specific differences in cuticular hydrocarbons of fig wasps, using the pollinator C. fusciceps as a case study.

Studies On Endocrine And Behavioral Assessment Of Reproductive Status In Asian Elephants (Elephas Maximus)

2013-01-16T07:33:41Z

Title: Studies On Endocrine And Behavioral Assessment Of Reproductive Status In Asian Elephants (Elephas Maximus) Authors: Ghosal, Ratna Abstract: The Asian elephant (Elephas maximus), a charismatic ‘flagship species’, is threatened by extinction in the wild, and the development of self-sustainable captive populations is a key conservation challenge. A third of the Asian elephant population is presently in captivity and information on the reproductive status, especially in females, is still lacking to a large extent. The onset of estrus in female Asian elephants is not associated with any visible physical signs, thus making the assessment of the reproductive status rather difficult. One approach to understanding reproductive cyclicity of animals is through generating profiles of reproductive hormones in blood (Wiseman et al. 1983; Brown et al. 1999, 2004). Profiles of reproductive hormones such as progesterone (P4), estrogen, luteinizing hormone and follicle stimulating hormone have already being demonstrated in Asian elephants (Brown et al. 1999, 2004; Brown 2000). In these studies, the reproductive status of females, maintained in zoos/captivity, was characterized based on circulating levels of hormones in blood samples. This is difficult to implement in the case of semi-captive or wild populations of elephants due to practical, legal and ethical considerations. In order to overcome this problem and to better understand the estrous status of female elephants, it is important to develop and validate non-invasive methods to monitor the reproductive status of female Asian elephants. An alternative approach to evaluating the reproductive status of females is to consider behavioral responses shown by males towards chemical signals produced by females to advertise their reproductive status. In order to understand the reproductive status of the individual belonging to the opposite sex, studies have shown that elephants rely on a variety of chemical signals produced in biological fluids such as urine, temporal gland secretion, inter-digital gland secretion, etc. (Krishnan 1972; Rasmussen & Schulte 1998). Chemical signaling is one of the prominent modes of communication in elephants, especially with respect to locating potential mates (Sukumar 2003). Thus, in most cases, elephants usually employ specific behavioral responses, for example sniff, check and place behaviors of trunk, for investigating the reproductive status of the conspecific individual, belonging either to the same or the opposite sex (Rasmussen et al. 1996; Schulte & Rasmussen 1999; Bagley et al. 2006). The objectives of the thesis are two-fold. First, to develop a non-invasive method of reproductive monitoring from fecal hormonal metabolites and also to understand the possible role of feces as an inter-sexual signal. The main body of thesis is divided into four chapters. 1) Development and validation of a non-invasive method to estimate progesterone metabolite in feces, to monitor the reproductive cyclicity of female elephants (Chapter 2). 2)Generation and characterization of progesterone and its metabolite, 5α-P-3-OH, profiles of semi-captive females using the developed non-invasive method to measure fecal metabolites (Chapter 3). 3) Validation of developed methodology and assay systems to a wild-population of female elephants (Chapter 4). 4) Feces as a potential source for inter-sexual chemical signaling in Asian elephants (Chapter 5). The above studies were carried out on semi-captive male and female elephants maintained in the forest camps of Mudumalai Wildlife Sanctuary (MWLS), Tamil Nadu and Bandipur National Park (BNP), Karnataka, India (Chapters 2, 3 and 5). For Chapter-4, free-ranging females of the MWLS were examined. 1. Development and validation of a non-invasive method to estimate progesterone metabolite in feces, to monitor the reproductive cyclicity of female elephants Niemuller et al. (1993) generated a profile of the progesterone metabolite, 5βpregnanetriol, to assess the estrous phase of Asian elephants based on non-invasive urine sampling. However, the collection of urine is difficult and to some extent impossible in the case of semi-captive and as well as that of wild elephants. Thus, the method of choice in this study was the development and validation of a non-invasive approach to measure fecal progesterone metabolites to assess reproductive status of females. Sampling was carried out at monthly intervals on three female elephants at the MWLS forest camp, while three other females maintained at the BNP forest camp were sampled fortnightly. An enzyme linked immuno-sorbent assay was developed to measure the concentration of the progesterone metabolite, 5α-P-3OH in the fecal samples of the semi-captive females. Using varying concentrations of the hapten (5α-P3OH), from low (0.1 mM) to high (1 mM), a standard curve was first generated, which had a linear range between 0.25 mM to 62.5 mM, with an EC50 of 1.37 mM. The linear range was then used to detect the concentrations of 5α-P-3OH in the fecal samples of females examined. The non-invasive method was further validated as there existed a positive correlation (p<0.1) between the levels of fecal 5α-P-3OH and that of concentration of circulating P4, measured in blood samples. This is the first valid documentation of a non-invasive method based on fecal progesterone metabolite pattern in order to assess the reproductive status of the female Asian elephants. 2. Generation and characterization of reproductive hormone profiles of semi-captive females using the developed non-invasive method to measure fecal metabolites With the establishment of a non-invasive method to understand occurrence of estrus in female elephants (described in Chapter 2), attempts were made to generate hormonal profiles over a longer time interval through more frequent sampling. Based on sampling at weekly intervals, the concentrations of both fecal 5α-P-3OH and that of native P4 hormone in the blood were determined, and the females were then identified as belonging to different reproductive states of hormonal cycling (follicular and luteal phases), non-cycling and pregnant categories. Of the 7 females sampled at both MWLS and BNP, three distinct categories of hormone-metabolite profiles emerged. The first category included four females that showed regular cyclicity throughout the sampling period, as monitored through the measurement of fecal 5α-P-3OH and serum P4 levels. The estrous cycle of all the four females was divided into two phases (follicular and luteal), based on the patterns of fecal 5α-P-3OH and serum P4 concentrations. The follicular phase of the estrous cycle was assigned when the values of both fecal 5α-P-3OH and serum P4 remained below 0.3 µg/gm and 0.3 ng/ml, respectively, for a considerable time period (viz. >2 wk). However, the luteal phase was characterized, when the levels of fecal 5α-P-3OH and serum P4 remained at or above 0.3 µg/gm and 0.3 ng/ml respectively, over a period of more than 2 - 3 wk. The second category had two females showing a ‘flat-lining pattern’ for the levels of fecal 5α-P-3OH and serum P4 concentrations, without any peak or dip in their concentrations. Since the reproductive pattern for both these females was flat-lined throughout their sampling period (51 wk), maintaining the levels of fecal 5α-P-3OH and serum P4 below 0.3 µg/gm and 0.3 ng/ml respectively, the females were considered to be non-cycling or anestrus. The third category had one female in which the levels of both fecal 5α-P-3OH and serum P4 measured were consistently high throughout the entire sampling period (26 wk). The levels of both fecal -P-3OH and serum P4 were above 0.3 µg/gm and 0.3 ng/ml, respectively. At the end of the sampling period, this female delivered a male calf; thus, the measured concentrations of fecal 5α-P-3OH and serum P4 signified the levels maintained during the gestational phase of this female. Sampling and hormonal analyses were also carried out for a male in the MWLS forest camps to investigate the baseline concentrations of fecal 5α-P-3OH and serum P4. The male showed consistently low concentrations of both fecal 5α-P-3OH and serum P4 throughout the sampling period (16 wk), below the margin of 0.3 µg/gm and 0.3 ng/ml, respectively. In this part of the study, it was confirmed that the reproductive status of a female elephant can be correctly assessed on the basis of measurements of fecal 5α-P-3OH alone, with repeated sampling of the female over a longer time scale. It was also shown that the strength of the positive correlation between the concentrations of the fecal 5α-P3OH and the serum P4 increased (p<0.01) for a larger sample size than that obtained for a comparatively smaller sample (described in Chapter 2). 3. Validation of developed methodology and assay systems to wild-population of female elephants So far, the non-invasive method to estimate fecal progesterone metabolite in order to predict occurrence of estrus of elephants was largely applied to semi-captive females (described in Chapters 2, 3). However, the necessity of such a method is being recognized for assessing the reproductive status of free-ranging females. Several findings have described differences in the rate and type of steroid metabolite excretion among individuals maintained under different diet regimes (Wasser et al. 1993; Smith et al. 2006). For instance, female elephants in the forest camps are provided with supplementary diet consisting of sugarcane, rice and millets. This diet is strikingly different from the feeding materials consumed by wild/free-ranging elephants (Sukumar 2003). Therefore, differences in dietary components can potentially influence the fecal steroid metabolites’ excretory patterns shown by semi-captive vis-a-vis wild females, which can affect the validity of measuring fecal 5α-P-3OH to predict females’ reproductive status. In order to examine this problem, the non-invasive method was applied to the population of wild elephants in the forests of MWLS, through random one time sampling of 30 individual female elephants. The steroid extraction efficiency (73 ± 11.0%, mean ± S.D., n =30) determined for the fecal samples collected from the wild females was not significantly different from the coefficient calculated in the case of the semi-captive females (80 ± 4.3%, mean ± S.D., n = 38). This indicated that dietary differences between wild and captive elephants did not influence levels of fecal hormonal metabolites in feces, unlike earlier observations on baboons (Wasser et al. 1993), old world primates (Wasser et al. 1988) and sheep (Smith et al. 2006). The values of the fecal allopregnanolone determined in the case of the wild females, ranged from as low as 0.06 µg/gm to as high as 23µg/gm of the sample, thus showing the heterogeneity of the samples, indicating that the females may be belonging to different reproductive phases. However, since sampling was carried out randomly, with an adult female being sampled just once, at this stage, it is not possible to identify or elaborate on the reproductive phase of the females. This is the first study reporting the values of the fecal progesterone metabolite in female Asian elephants in the wild. Further studies may be required to carry out long term monitoring of the wild females, through repeated collection of fecal samples over time from particular female. 4. Feces as a potential source for inter-sexual chemical signaling in Asian elephants In this Chapter-5, behavioral trials were conducted on male elephants to understand the role of fecal matter in conveying inter-sexual chemical signals. This was demonstrated by analyses of specific chemosensory behavioral responses shown by males towards the fecal samples of females that were strangers and belonging to different reproductive stages. Males showed four prominent behavioral responses namely ‘distant sniff’, ‘close sniff’, ‘check’ and ‘place’ towards the fecal samples of females. The sum of frequencies of these four responses (distant sniff, close sniff, check and place) was much higher for samples of the follicular (pre-ovulatory) phase females as compared to those of the luteal (post-ovulatory) phase females (p<0.005). Thus, for the first time, it was shown that male elephants are able to discriminate the different reproductive phases of females based on their specific behavioral response towards the fecal samples of the opposite sex. In conclusion, the thesis has focused on understanding and providing new insights regarding the reproductive biology of the female Asian elephants. This has been achieved through the development of the non-invasive method based on measuring the concentrations of the fecal progesterone metabolite and through the analyses of the chemosensory responses performed by the males towards the fecal samples of strange females. These methods can potentially be applied to the populations of both wild and captive/semi-captive female elephants in order to evaluate their reproductive status, through non-invasive measures. The information derived from the application of such methods will help in understanding the reproductive potential of the wild elephants under various environmental and ecological conditions. Further, the non-invasive measurement of reproductive hormones will help in monitoring the reproductive state of the individuals and thus aid in planning strategies for the welfare and management of the elephants maintained in captive or semi-captive conditions.