Stuart Parsons

Echolocation call intensity in the aerial hawking bat Eptesicus bottae (Vespertilionidae) studied using stereo videogrammetry.

SUMMARY Aerial hawking bats use intense echolocation calls to search for insect prey. Their calls have evolved into the most intense airborne animal vocalisations. Yet our knowledge about call intensities in the field is restricted to a small number of species. We describe a novel stereo videogrammetry method used to study flight and echolocation behaviour, and to measure call source levels of the aerial hawking bat Eptesicus bottae (Vespertilionidae). Bats flew close to their predicted minimum power speed. Source level increased with call duration; the loudest call of E. bottae was at 133 dB peSPL. The calculated maximum detection distance for large flying objects (e.g. large prey, conspecifics) was up to 21 m. The corresponding maximum echo delay is almost exactly the duration of one wing beat in E. bottae and this also is its preferred pulse interval. These results, obtained by using videogrammetry to track bats in the field, corroborate earlier findings from other species from acoustic tracking methods.

Kiwi forego vision in the guidance of their nocturnal activities.

BACKGROUND In vision, there is a trade-off between sensitivity and resolution, and any eye which maximises information gain at low light levels needs to be large. This imposes exacting constraints upon vision in nocturnal flying birds. Eyes are essentially heavy, fluid-filled chambers, and in flying birds their increased size is countered by selection for both reduced body mass and the distribution of mass towards the body core. Freed from these mass constraints, it would be predicted that in flightless birds nocturnality should favour the evolution of large eyes and reliance upon visual cues for the guidance of activity. METHODOLOGY/PRINCIPAL FINDINGS We show that in Kiwi (Apterygidae), flightlessness and nocturnality have, in fact, resulted in the opposite outcome. Kiwi show minimal reliance upon vision indicated by eye structure, visual field topography, and brain structures, and increased reliance upon tactile and olfactory information. CONCLUSIONS/SIGNIFICANCE This lack of reliance upon vision and increased reliance upon tactile and olfactory information in Kiwi is markedly similar to the situation in nocturnal mammals that exploit the forest floor. That Kiwi and mammals evolved to exploit these habitats quite independently provides evidence for convergent evolution in their sensory capacities that are tuned to a common set of perceptual challenges found in forest floor habitats at night and which cannot be met by the vertebrate visual system. We propose that the Kiwi visual system has undergone adaptive regressive evolution driven by the trade-off between the relatively low rate of gain of visual information that is possible at low light levels, and the metabolic costs of extracting that information.

Advantages and Disadvantages of Techniques for Transforming and Analyzing Chiropteran Echolocation Calls

Abstract Bat researchers currently use a variety of techniques that transform echolocation calls into audible frequencies and allow the spectral content of a signal to be viewed and analyzed. All techniques have limitations and an understanding of how each works and the effect on the signal being analyzed are vital for correct interpretation. The 3 most commonly used techniques for transforming frequencies of a call are heterodyne, frequency division, and time expansion. Three techniques for viewing spectral content of a signal are zero-crossing, Fourier analysis, and instantaneous frequency analysis. It is important for bat researchers to be familiar with the advantages and disadvantages of each technique.

Terrestrial locomotion of the New Zealand short-tailed bat Mystacina tuberculata and the common vampire bat Desmodus rotundus.

SUMMARY Bats (Chiroptera) are generally awkward crawlers, but the common vampire bat (Desmodus rotundus) and the New Zealand short-tailed bat (Mystacina tuberculata) have independently evolved the ability to manoeuvre well on the ground. In this study we describe the kinematics of locomotion in both species, and the kinetics of locomotion in M. tuberculata. We sought to determine whether these bats move terrestrially the way other quadrupeds do, or whether they possess altogether different patterns of movement on the ground than are observed in quadrupeds that do not fly. Using high-speed video analyses of bats moving on a treadmill, we observed that both species possess symmetrical lateral-sequence gaits similar to the kinematically defined walks of a broad range of tetrapods. At high speeds, D. rotundus use an asymmetrical bounding gait that appears to converge on the bounding gaits of small terrestrial mammals, but with the roles of the forelimbs and hindlimbs reversed. This gait was not performed by M. tuberculata. Many animals that possess a single kinematic gait shift with increasing speed from a kinetic walk (where kinetic and potential energy of the centre of mass oscillate out of phase from each other) to a kinetic run (where they oscillate in phase). To determine whether the single kinematic gait of M. tuberculata meets the kinetic definition of a walk, a run, or a gait that functions as a walk at low speed and a run at high speed, we used force plates and high-speed video recordings to characterize the energetics of the centre of mass in that species. Although oscillations in kinetic and potential energy were of similar magnitudes, M. tuberculata did not use pendulum-like exchanges of energy between them to the extent that many other quadrupedal animals do, and did not transition from a kinetic walk to kinetic run with increasing speed. The gait of M. tuberculata is kinematically a walk, but kinetically run-like at all speeds.

Echolocation calls and wing morphology of bats from the West Indies

Echolocation calls of 119 bats belonging to 12 species in three families from Antillean islands of Puerto Rico, Dominica, and St. Vincent were recorded by using time-expansion methods. Spectrograms of calls and descriptive statistics of five temporal and frequency variables measured from calls are presented. The echolocation calls of many of these species, particularly those in the family Phyllostomidae, have not been described previously. The wing morphology of each taxon is described and related to the structure of its echolocation calls and its foraging ecology. Of slow aerial-hawking insectivores, the Mormoopidae and Natalidae Mormoops blainvillii, Pteronotus davyi davyi, P. quadridens fuliginosus, and Natalus stramineus stramineus can forage with great manoeuvrability in background-cluttered space (close to vegetation), and are able to hover. Pteronotus parnellii portoricensis is able to fly and echolocate in highly-cluttered space (dense vegetation). Among frugivores, nectarivores and omnivores in the family Phyllostomidae, Brachyphylla cavernarum intermedia is adapted to foraging in the edges of vegetation in background-cluttered space, while Erophylla bombifrons bombifrons, Glossophaga longirostris rostrata, Artibeus jamaicensis jamaicensis, A. jamaicensis schwartzi and Stenoderma rufum darioi are adapted to foraging under canopies in highly-cluttered space and do not have speed or efficiency in commuting flight. In contrast, Monophyllus plethodon luciae, Sturnira lilium angeli and S. lilium paulsoni are adapted to fly in highly-cluttered space, but can also fly fast and efficiently in open areas.

PHYLOGENETICS OF SMALL HORSESHOE BATS FROM EAST ASIA BASED ON MITOCHONDRIAL DNA SEQUENCE VARIATION

Abstract We undertook analyses of mitochondrial DNA gene sequences and echolocation calls to resolve phylogenetic relationships among the related bat taxa Rhinolophus pusillus (sampled across China), R. monoceros (Taiwan), R. cornutus (main islands of Japan), and R. c. pumilus (Okinawa, Japan). Phylogenetic trees and genetic divergence analyses were constructed by combining new complete mitochondrial cytochrome-b gene sequences and partial mitochondrial control region sequences with published sequences. Our work showed that these 4 taxa formed monophyletic groups in the phylogenetic tree. However, low levels of sequence divergence among the taxa, together with similarities in body size and overlapping echolocation call frequencies, point to a lack of taxonomic distinctiveness. We therefore suggest that these taxa are better considered as geographical subspecies rather than distinct species, although this should not diminish the conservation importance of these island populations, which are important evolutionarily significant units. Based on our findings, we suggest that the similarities in body size and echolocation call frequency in these rhinolophids result from their recent common ancestry, whereas similarities in body size and call frequency with R. hipposideros of Europe are the result of convergent evolution.

Identification of New Zealand bats (Chalinolobus tuberculatus and Mystacina tuberculata) in flight from analysis of echolocation calls by artificial neural networks

Time-expanded and heterodyned echolocation calls of the New Zealand long-tailed Chalinolobus tuberculatus and lesser short-tailed bat Mystacina tuberculata were recorded and digitally analysed. Temporal and spectral parameters were measured from time-expanded calls and power spectra generated for both time-expanded and heterodyned calls. Artificial neural networks were trained to classify the calls of both species using temporal and spectral parameters and power spectra as input data. Networks were then tested using data not previously seen. Calls could be unambiguously identified using parameters and power spectra from time-expanded calls. A neural network, trained and tested using power spectra of calls from both species recorded using a heterodyne detector set to 40 kHz (the frequency with the most energy of the fundamental of C. tuberculatus call), could identify 99% and 84% of calls of C. tuberculatus and M. tuberculata, respectively. A second network, trained and tested using power spectra of calls from both species recorded using a heterodyne detector set to 27 kHz (the frequency with the most energy of the fundamental of M. tuberculata call), could identify 34% and 100% of calls of C. tuberculatus and M. tuberculata, respectively. This study represents the first use of neural networks for the identification of bats from their echolocation calls. It is also the first study to use power spectra of time-expanded and heterodyned calls for identification of chiropteran species. The ability of neural networks to identify bats from their echolocation calls is discussed, as is the ecology of both species in relation to the design of their echolocation calls.

Classification of Echolocation Calls from 14 Species of Bat by Support Vector Machines and Ensembles of Neural Networks

Calls from 14 species of bat were classified to genus and species using discriminant function analysis (DFA), support vector machines (SVM) and ensembles of neural networks (ENN). Both SVMs and ENNs outperformed DFA for every species while ENNs (mean identification rate – 97%) consistently outperformed SVMs (mean identification rate – 87%). Correct classification rates produced by the ENNs varied from 91% to 100%; calls from six species were correctly identified with 100% accuracy. Calls from the five species of Myotis, a genus whose species are considered difficult to distinguish acoustically, had correct identification rates that varied from 91 – 100%. Five parameters were most important for classifying calls correctly while seven others contributed little to classification performance.

Evolution of Brain Size in the Palaeognath Lineage, with an Emphasis on New Zealand Ratites

Brain size in vertebrates varies principally with body size. Although many studies have examined the variation of brain size in birds, there is little information on Palaeognaths, which include the ratite lineage of kiwi, emu, ostrich and extinct moa, as well as the tinamous. Therefore, we set out to determine to what extent the evolution of brain size in Palaeognaths parallels that of other birds, i.e., Neognaths, by analyzing the variation in the relative sizes of the brain and cerebral hemispheres of several species of ratites and tinamous. Our results indicate that the Palaeognaths possess relatively smaller brains and cerebral hemispheres than the Neognaths, with the exception of the kiwi radiation (Apteryx spp.). The external morphology and relatively large size of the brain of Apteryx, as well as the relatively large size of its telencephalon, contrast with other Palaeognaths, including two species of historically sympatric moa, suggesting that unique selective pressures towards increasing brain size accompanied the evolution of kiwi. Indeed, the size of the cerebral hemispheres with respect to total brain size of kiwi is rivaled only by a handful of parrots and songbirds, despite a lack of evidence of any advanced behavioral/cognitive abilities such as those reported for parrots and crows. In addition, the enlargement in brain and telencephalon size of the kiwi occurs despite the fact that this is a precocial bird. These findings form an exception to, and hence challenge, the current rules that govern changes in relative brain size in birds.

Echolocation Calls of the Long-Tailed Bat: A Quantitative Analysis of Types of Calls

The echolocation calls of long-tailed bats (Chalinolobus tuberculatus) were recorded in the Eglinton Valley, Fiordland, New Zealand, and digitized for analysis with signal-processing software. Univariate and multivariate analyses of measured features facilitated a quantitative classification of the calls. Cluster analysis was used to categorize calls into two groups equating to search and terminal-buzz calls described qualitatively for other species. When moving from search to terminal phases, the calls decrease in bandwidth, maximum and minimum frequency of call, and duration. Search calls begin with a steep-downward FM sweep followed by a short, less-modulated component. Buzz calls are FM sweeps. Although not found quantitatively, a broadband pre-buzz group of calls also was identified. Ambiguity analysis of calls from the three groups shows that search-phase calls are well suited to resolving the velocity of targets, and hence, identifying moving targets in stationary clutter. Pre-buzz and buzz calls are better suited to resolving range, a feature that may aid the bats in capture of evasive prey after it has been identified.