Call Diversity, Spatio-Temporal Patterning And Masking Interference In An Assemblage Of Acoustically Communicating Ensiferan Species Of A Tropical Evergreen Forest In Southern India
The present study provides the first description of the calls of a multi-species ensiferan assemblage in a tropical evergreen forest of the Indian subcontinent. I have identified and described the calls of twenty ensiferan species constituting the nocturnal acoustic community of a tropical evergreen forest in KNP. I found that the multi-species ensiferan assemblage consisted of diverse taxa representing subfamilies of the families Gryllidae, Tettigoniidae and Anostostomatidae. Eight acoustically communicating species of the family Gryllidae were found. Two species belonged to the family Mogoplistidae. Interestingly, each subfamily was mostly represented by only one genus. In the tettigoniids, representative species were found only from subfamilies Pseudophyllinae, Phaneropterinae and Mecopodinae. The species richness of the acoustically communicating cricket assemblage in the tropical evergreen forest of Kudremukh was found to be low. This study did not include non-calling and ultrasonic species of crickets. The gryllid and tettigoniid species analyzed exhibited different frequency (both narrow and broadband) and temporal patterns. Species belonging to the family Gryllidae had narrow band calls (with bandwidths not greater than 1 kHz) and had dominant frequencies mainly between 3 and 7 kHz. The calls of tettigoniids covered a wide spectral range reaching far into the ultrasound in species of the genus Mecopoda. Interestingly, of nine tettigoniid species, the calls of four (Onomarchus sp., Phyllomimus sp., Brochopeplus sp. and ‘15 kHz’) were narrow band and in the audible range, similar to those of gryllids. Although there was a high overlap of call frequencies between 3 to 7 kHz, gryllid species separated in their syllable repetition rate, which varied from six syllables per second in Landreva to 60 syllables/second in Gryllitara. Species with overlapping syllable repetition rates of 10 – 20 syllables per second separated along the frequency axis. There were species such as those of Phaloria and Gryllitara, Scapsipedus, Xabea and Callogryllus that overlapped both in the spectral and syllable repetition rates. These species however, differed in the other temporal features such as call duration, call period and number of syllables per call. This study also provides the first description of the calls and stridulatory structures of an Indian weta species (Family Anostostomatidae). Both males and females of this species were found to stridulate. The calls of the two sexes had similar spectral features. Male calls consisted of four syllables each, while female calls were bisyllabic. Stridulatory structures were similar between the sexes. I also quantitatively validated the reliability of human listener - based psychoacoustic sampling as a technique to monitor species richness and relative abundance of acoustically communicating ensiferan species that are within the human hearing range. I have shown using controlled psychoacoustic tests in the laboratory that a trained listener is capable of identifying the species as well as the number of individuals of Ensifera with high accuracy. This study suggests that trained listener - based psychoacoustic sampling may be preferable to carry out rapid assessments and species inventories of gryllids and low frequency katydid species in tropical forests. My study also suggests that acoustic monitoring of Orthoptera should be done using both the trained listener - based spot sampling and ambient noise recordings using ultrasound detectors for accurately estimating species richness and relative abundance in an area. Using focal animal sampling, I have shown that most species in the tropical forest ensiferan assemblage of Kudremukh National Park did not move more than a metre in a span of half an hour. The acoustic sampling should be designed in such a way as to cause minimal disturbance to the calling animals and could be limited to ten minutes to avoid re-counting individuals and counter the problem of pseudoreplication. I also investigated the spatial dispersion of calling sites in the vertical dimension. This study revealed vertical stratification of the calling heights of the twenty ensiferan species. Calling heights of both gryllid and tettigoniid species ranged from the ground to the canopy, although more gryllid than tettigoniid species occupied the ground and herb layer. Post hoc comparisons and cluster analysis indicated the presence of discrete calling height layers corresponding to the canopy, understorey, herb and ground layer. These clusters emerged from the raw data of calling heights of individuals of each species without a priori distinction of layers. This is in contrast to other studies on vertical stratification in arthropods and bats where baits, traps and mist nets are placed at different vertical layers, thereby demarcating the layers beforehand. Previous studies on crickets, cicadas and frogs have shown preference for the height of calling sites qualitatively. To my knowledge, this is the first study to quantitatively establish vertical stratification in calling heights in an ensiferan assemblage of an evergreen forest. No correlation between the calling heights and mean dominant frequencies of the species were found. Cricket species with relatively low frequency calls (3–4 kHz) occupied both the ground layer (Callogryllus sp. and Scapsipedus sp.) and the canopy (Xabea sp. and Onomarchus sp) suggesting that these narrow-band, relatively low frequency signals may be optimal for sound transmission in the cluttered habitat of the forest floor (due to leaf litter) and the canopy (due to high leaf density). Species with high frequencies such as Brochopeplus sp. and ‘15 kHz’ called mainly from vegetation in the understorey. Species with broadband calls (Mecopoda sp., Pirmeda sp. and Elimaea sp.) called just above the ground layer and from the understorey suggesting that calls with higher frequencies and bandwidths may be used in the somewhat less-cluttered microhabitat of the understorey. Calling height stratification in the ensiferan assemblages of tropical forests could also be due to other ecological factors such as predation by spiders, mantises, bats, birds or primates. The wide range of duty cycles, presence of high duty cycle callers (such as Mecopoda) and the lack of correlation of duty cycle with calling height found in our study site are interesting. Studies on acoustic transmission in different microhabitats at different heights and on predation pressure on the ensiferan species will provide further insight into the selective forces influencing calling height stratification. The multi-species assemblage constituting the nocturnal acoustic community was found to be calling in the same time period between evening to midnight and no species was found to have a unique calling time that is different from that of another species. There was no diel partitioning of calling time between the acoustically communicating ensiferan species. Frogs and cicadas that can be considered as acoustic competitors of the ensiferan assemblage appeared to be separating from crickets on a seasonal and diel scale respectively. This study has quantified the amount of masking interference in three dimensions viz. temporal, fine temporal and spectral, between sixteen species belonging to the nocturnal acoustic ensiferan assemblage of an evergreen forest. Frequency histograms of overlap, bar graphs of overlap on a species by other species and Mantel’s test results on matrix correlation suggest negative relations between the temporal, fine-temporal and spectral overlaps. Species with high overlap in one dimension had very low levels of overlap in any of the other two dimensions, suggesting acoustic resource partitioning in the ensiferan assemblage of the evergreen forest. I also tried to quantify the extent of spatial overlap between species based on calling intensity and inter-specific distances. However, spatial overlap could not be analysed further as there were some species pairs for which I did not have the inter-individual distances despite carrying out the field work for six months. The procedure of estimating spatial overlap between species pairs and the result along with missing gaps is presented in appendix 2. It will be interesting to investigate the extent of spatial overlap between species pairs as the fourth dimension in which species could separate to avoid acoustic competition. It is also important to estimate the relative abundance of species in the evergreen forest to obtain a realistic representation of masking interference between species. Partitioning of acoustic resources among ensiferan assemblage could also be better explained by analysing all the dimensions.
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