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Applying fixed target strengths (TS) to individual insects when modelling the detection distances of prey by bats echolocating at different frequencies, is fundamentally flawed.

Posted by Leonard on 19 May 2008 at 22:50 GMT

We estimated detection distances with a standardized set of three prey types: a small insect (mealworm head on), as well as a small and a large moth with target strengths of −40 dB, −20 dB, and −5 dB, respectively [25], [26]
http://plosone.org/article/info:doi/10.1371/journal.pone.0002036#article1.body1.sec2.sec4.p1

Introduction.
In their brief explanation of the method used to calculate the detection distances of differently-sized insect prey, the authors of this paper have made a conceptual error which has led to incorrectly calculated results. This in turn, may have a bearing on the interpretation of the results and the conclusions drawn from them in the paper’s discussion.

Critique.
The authors evidently intended to calculate the detection distances of three size categories of insect prey for bats echolocating at different call intensities and different echolocation frequencies. However, rather than defining the three categories of insects using their size in terms of maximum planar cross-sectional area (CSA), the authors have chosen to categorize them in terms of target strength (TS). This is conceptually wrong as the TS of prey is affected by Rayleigh scattering in addition to prey CSA and is therefore, dependent on the frequency of ensonification. This is clearly seen in Fig. 1, which is a theoretical equivalent of the empirical data provided in Fig. 2a by Surlykke et al. (1999), (authors’ ref [25]). The empirical data in the latter reference evidently contains some degree of offset and experimental error.

Please view Figure 1 and Tables 1a, b, and c here: http://www.plosone.org/attachments/pone.0002036.kerry.pdf

From Fig. 1, it is clearly seen that if a constant TS line is drawn horizontally from the -20dB point on the y-axis, the equivalent CSA on the x-axis is different for each of the three frequency curves depicted. From this, we can conclude that the prey within each of the three TS categories proposed by the authors for calculating prey detection distances, are non-identical at each frequency (and for each bat). This is contrary to the intention of the authors as they specifically name one of the prey as being the ‘front-on’ cross-sectional-area of a mealworm, specifically indicating that the CSA of this insect (and by inference, that of the other two insects) was meant to be identical for each bat (and frequency) in the detection distance calculations. Fig. 1 clearly indicates that the sizes of prey are not consistent when identical TS’s are used for each frequency in the detection distance calculations.

In table 1b, I have calculated the effective CSA of the three prey TS’s (-40, -20 and -5dB) at each ensonification frequency, summarized in Table 1a, as specified by the author. Not only does Table 1b show the different effective CSA’s (and sizes) of the ‘same’ insect at different echolocation frequencies due to the effects of Rayleigh scattering, it also shows that the CSA of a mealworm only approaches a realistic ‘front-on’ CSA for this insect at higher ensonification frequencies. Although not specified by the authors, I additionally calculated the CSA of the mealworm at 100 kHz, which turned out to be 8.6 mm2. Assuming the ‘end-on’ CSA of the mealworm to be approximately circular in form, this equates to a mealworm diameter of 3.3mm, which approximates the diameter of mealworms with which I am familiar! For this reason, I have assumed for the calculations in Table 1c that the three TS categories used by the authors were obtained from three sizes (CSA) of prey at 100 kHz (although it is possible at a pinch that the authors used the TS values calculated at their highest reporting frequency of 70 kHz).

Detection Distance Modelling.
Table 1c presents the results of detection distance calculations based on the CSA of three sizes of insect prey, namely a mealworm with CSA of 8.6 mm2, a small moth of 105 mm2 and a large moth of 3162 mm2. The TS of each of these insects is different at each frequency due to their individual CSA’s and the frequency-dependent effects of Rayleigh scattering on target echoes. This is particularly evident in the detection distances calculated for the mealworm prey. Note that several of the detection distances calculated for the small mealworm CSA are very small, which may make them effectively invisible to some echolocating bats due to pulse-echo overlap. On the other hand, the Ts of the largest prey is similar for at all specified frequencies due to the reduced effects of Rayleigh scattering on large CSA prey.

The model used for calculating the detection distances in Table 1c differs from that typically used by others in that it does not treat planar targets as if they were spherical, which typically leads to over-estimation of distance for small and short-range targets. The model also accounts for the aperture of bat pinnae, which avoids additional over-estimation of distances for small and short-range targets. A conservatively sized pinna aperture (or CSA) of 113 mm2 was assumed for all species of bats addressed by this paper.

Conclusion.
The concept of using fixed values of TS instead of fixed values of CSA for any insect prey at different frequencies is erroneous as it ignores the effects of Rayleigh scattering and is equivalent to changing the size (CSA) of the ‘same’ insect for each bat echolocating at a different frequency. Consequently, the methodology employed by the authors for calculating detection distances of prey in this paper requires significant revision.

Some of the re-calculated detection distances using fixed values of CSA for the three categories of prey (and particularly for the mealworm), differ significantly from those calculated by the authors using fixed TS. Consequently, some of the statements made and conclusions drawn by the authors in their discussion may require revision.

RE: Applying fixed target strengths (TS) to individual insects when modelling the detection distances of prey by bats echolocating at different frequencies, is fundamentally flawed.

surlykke replied to Leonard on 23 May 2008 at 07:35 GMT

We estimated detection range for prey to illustrate the influence of distance and atmospheric attenuation at the different frequencies employed by the bats in our study. The two target strengths for moths are measured in Surlykke et al. -99, where target strengths did not depend much on frequency at frequencies above 10 kHz. No theoretical model can account for the complicated shape and surface of an insect. Even our measured values are only indications of the target strengths, since a live flying insect will reflect a much more variable echo, as shown already by Roeder and later also others. The reason why we included TS for the (artificial prey) mealworm head-on, was to see what happened at very short distances, but you are right: Here frequency vs. size would play a significant role, and in retrospect, mealworms might have been omitted. However, our results and discussion are based on the two moth sizes, and the main conclusion holds: the detection distances for small and big moths are far more similar in this assemblage of bats, than the variation in output intensity would suggest at first glance, due to the link between high intensity and high frequency. Thus, the data indicate that insectivorous bats operating in open/edge space need a minimum detection range and thus turn up their volume if the absorption is too severe.
Annemarie Surlykke and Elisabeth Kalko.

RE:Applying fixed target strengths (TS) to individual insects when modelling the detection distances of prey by bats echolocating at different frequencies, is fundamentally flawed.

Leonard replied to surlykke on 02 Jun 2008 at 01:33 GMT

Although not the subject of this discussion, it should be pointed out that the theoretical target strengths presented in figure 1 of the Critique do not represent the target strengths of volant prey as suggested by the authors. Figure 1 of the Critique is the result of modelling stationary moths with outspread wings to obtain the maximum cross-sectional area, which is equivalent to the method used by Surlykke et al. (1999) to obtain their empirical data.

Contrary to the view of the authors in their response, figure 2a of Surlykke et al. (1999) and figure 1 of the Critique clearly show that the target strengths of prey with cross-sectional areas between 100 and 1000mm2 vary significantly with frequencies both below and above 10 kHz.

From their response, it is clear that the failure of the authors to take account of Rayleigh scattering as a frequency-dependent component of target strength estimates (at least for the hypothetical mealworm-sized prey), is a consequence of:-

• Assuming linear regression for the three frequency curves in fig. 2a of Surlykke et al. (1999) and consequently, assumed linear relationships between TS and CSA.
• Assuming identity of the 30 kHz and 100 kHz curves in fig. 2a of Surlykke et al. (1999), even when the data correctly suggests a tendency towards different gradients.
• Extrapolating the assumed single linear regression line representing both 30 kHz and 100 kHz beyond the lowest CSA data point in fig. 2a of Surlykke et al. (1999), by a factor of 23 to obtain the target strength of mealworm-sized prey.

Given apparent agreement on the inaccuracy of the detection distance estimates for mealworm-sized prey provided by the authors, a revision of table 2 in the web version of the paper is in order to avoid the use and quotation of invalid detection distance data by future authors.

The detection distance data provided in Table 1c of the Critique, which accounts for the effects of Rayleigh scattering on echoes, suggests the following sections of text also need revision:

Detection ranges; Para. 2
For small targets, the maximum detection ranges of the two Noctilio species, which emitted the loudest calls exceeding all other intensity data known so far from free-flying bats, were still within the general range of all other study species.

Detection ranges; Para. 3
Comparison of maximum detection ranges suggests that detection probabilities of small insects are indeed rather similar across bats.

Detection ranges; Para. 6
Although maximum source levels varied by more than a factor of 10 the detection distances for all target strengths only varied by a factor 2–3 or less.

It would also be appropriate for some other parts of the text to undergo revision that refer to fixed target strengths instead of moth cross-sectional area when discussing methods.

Finally, whereas the data provided in figure 2a of Surlykke et al. (1999) relates specifically to Danish moths, comparable data has not been provided by the authors for the sizes of Panamanian moths. This omission is significant, as the sizes of moths available to and consumed by the bats of Panama ought to have influenced the choice of ‘moth sizes’ employed by the authors to ‘illustrate differences in prey detection distances. The differences in detection distances are also influenced by the sizes of moths associated with the terms ‘large’ and ‘small’ by the authors for the comparison.

Leonard Kerry.