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Bat Echolocation Researc h - Bat Conservation International

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Figure 5: Call sequence showing the frequency shift between two individuals of the same species (Pteronotus davyi) foraging in the same space.<br />

variation for key call parameters such as the maximum,<br />

minimum, and characteristic frequencies (FMIN, FMAX,<br />

FC), as well as pulse duration (DUR).<br />

Without a large series of recordings for even the most<br />

common species, I would not have observed the frequency<br />

shift in calls of individuals. This was noted by<br />

watching calls on the computer screen in real time with<br />

the Anabat system while observing the bats with a spotlight.<br />

Frequently an individual shifts the characteristic<br />

frequency up or down when another conspecific arrives<br />

in the same area. This shift generally ranges from 2-3<br />

kHz. An example of this frequency shift between two<br />

individuals of the same species (P. davyi) foraging in the<br />

same space can be seen in Fig. 5. Observers unaware of<br />

such frequency shifts when conspecifics use the same<br />

space could overestimate the variation in the basic<br />

search-phase calls of an individual or species.<br />

To illustrate this point, I extracted data for call<br />

parameters from the recordings (Fig. 5). If only 15 pulses<br />

were recorded at the beginning of the call sequence,<br />

the range in frequencies would be FMAX=70.8 kHz,<br />

FMIN=58.3 kHz, and FC=67.6 kHz. If only the 11 pulses<br />

at the end of the call sequence were recorded, these<br />

parameters would be FMAX=68.2 kHz, FMIN=58.0 kHz,<br />

and FC=61.6 kHz. If this sequence were recorded using<br />

time expansion, which cannot display bat calls in real<br />

time (Pettersson, pers. comm.), researchers may erroneously<br />

assume that they were seeing variation in the<br />

search-phase calls of one individual.<br />

Like any new endeavor, there must be a starting<br />

point. In the beginning, every call sequence recorded is<br />

by definition, an “unknown species” acoustically. It was<br />

clear there were many different types of calls based on<br />

shapes and frequency. The first task was to match faces<br />

with voices. This meant bats had to be captured for verification<br />

of species identification. Captured individuals<br />

were flown under controlled conditions, either in enclosures<br />

or in the field, to match recordings to positively<br />

identified animals. Chemiluminescent light tags were<br />

often used to allow released bats to be followed with the<br />

detector.<br />

At present, 28 of the 32 species of the non-phyllostomid<br />

bats (7 families) in Belize can be positively<br />

Section 2: Acoustic Inventories<br />

identified acoustically (Appendix 1). However, many<br />

other species that do not occur in Belize can also be<br />

identified acoustically. I have recorded and confirmed by<br />

capture dozens of common species that occur in Belize,<br />

Mexico, Honduras, Guatemala, El Salvador, Costa Rica,<br />

Venezuela, and Bolivia. I have not found geographic<br />

variation in the calls of these species to the extent that<br />

hinders identification after careful comparison to the<br />

Neotropical bat call libraries that are based on verified<br />

species calls.<br />

In the Neotropics, it can be both easy and difficult to<br />

identify bats using echolocation calls. It has been easy<br />

because with the multitude of vastly different families<br />

and genera, call sequences often separate readily. It has<br />

also been difficult because we do not fully understand<br />

the range of species distributions or, in many cases,<br />

species limits in the Neotropics.<br />

For example, there has been an increase in the<br />

number of recognized species since 1993 from 923 to an<br />

estimated 1,095 (Koopman 1993; Simmons, pers.<br />

comm.). This is an 18% increase. These taxonomic<br />

changes have implications for preconceived geographic<br />

ranges. Older published maps and museum identifications<br />

may no longer be valid (Simmons, pers. comm.).<br />

For several families (Emballonuridae, Noctilionidae,<br />

Mormoopidae), acoustic identification of species<br />

appears to be straightforward (O’Farrell and Miller 1997,<br />

1999). Others (e.g., Natalidae, Thyropteridae, Vespertilionidae,<br />

Molossidae) are more problematic. As my<br />

field techniques were refined and the limits of the equipment<br />

established, it was clear that acoustic identification<br />

in the field was not an option for species from two families.<br />

These were Natalidae, the funnel-eared bats, and<br />

Thyropteridae, the disc-winged bats.<br />

The Mexican funnel-eared bat (Natalus stramineus)<br />

appears to be a gleaner that is too quiet to detect in the<br />

field. This species is readily trapped and occasionally<br />

mist netted. A large number of fragmented recordings<br />

for a few individuals (n=24) were obtained, but only at<br />

very close range. Rydell et al. (2002) also found this to<br />

be the case when recording a single released individual<br />

in Mexico using a time-expansion system.<br />

Spix’s disc-winged bat (Thryoptera tricolor) is the only<br />

61

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