First published online November 19, 2007
Journal of Experimental Biology 210, 4104-4122 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.007930
Phylogenetic comparative analysis of electric communication signals in ghost knifefishes (Gymnotiformes: Apteronotidae)
Cameron R. Turner1,2,*,
Maksymilian Derylo3,4,
C. David de Santana5,6,
José A. Alves-Gomes5 and
G. Troy Smith1,2,7
1 Department of Biology, Indiana University, Bloomington, IN 47405,
USA
2 Center for the Integrative Study of Animal Behavior (CISAB), Indiana
University, Bloomington, IN 47405, USA
3 CISAB Research Experience for Undergraduates Program, Indiana University,
Bloomington, IN 47405, USA
4 Dominican University, River Forest, IL 60305, USA
5 Laboratório de Fisiologia Comportamental (LFC), Instituto Nacional
de Pesquisas da Amazônia (INPA), Manaus, AM 69083-000, Brazil
6 Smithsonian Institution, National Museum of Natural History, Division of
Fishes, Washington, DC 20560, USA
7 Program in Neuroscience, Indiana University, Bloomington, IN 47405,
USA

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Fig. 1. Signal parameter measurements. (A) EOD frequency trace (top) and
head-to-tail EOD waveform (voltage) trace (bottom). (B) Power spectrum (8192
points, Hanning window) of an EOD recording showing fundamental (F1), second
(F2) and third (F3) harmonic frequencies. (C) EOD frequency trace (top),
head-to-tail EOD waveform (voltage) trace (middle), and root-mean-square (RMS)
EOD amplitude trace (bottom) for a single chirp. Points used to measure signal
parameters are indicated with crosses and/or arrows. See
Table 1 for details on how
duration and FM parameters were calculated from these points. (D) EOD
frequency trace (top) and head-to-tail EOD waveform (voltage) trace (bottom)
for a chirp with extreme and prolonged reduction of EOD amplitude. Broken red
line indicates where EOD frequency was not measurable (i.e. when RMS EOD
amplitude dropped below 15% of baseline). Estimates of EODf were fixed at this
frequency until RMS EOD amplitude returned to at least 15% of its
baseline.
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Fig. 3. Photographs and species-typical electrocommunication signals of 15
apteronotid taxa. (A) Photograph of the head, not shown to scale. (B) A 3 ms
section of head-to-tail EOD waveform with peak-to-peak amplitude normalized
for all taxa. (C) EOD frequency trace (top) and head-to-tail EOD waveform
(voltage) trace (bottom) of characteristic chirps in each species. Amplitude
reductions during chirps are apparent from the amplitude envelope of the
waveform trace in some species. Axes are the same as in
Fig. 1A. Axis scales are
identical within column C across all panels to allow direct comparison of
chirp FM and duration. Red bars show positive start and stop times,
respectively. Orange bars show the negative stop time. (D) Additional examples
of chirps for each species shown as in C. Note the different frequency scale
in D compared with C. The frequency axis scales are identical across panels
within column D, but time scales vary as indicated. (E) Scatter plot of
positive FM and duration showing all EODMs recorded in each taxon. Chirps are
shown as red squares, GFRs as blue circles. The chirps identified in columns C
and D are colored yellow and labeled. The purple line illustrates the linear
function used to distinguish chirps from GFRs when necessary (see Materials
and methods for details). Note that the duration axis is logarithmic, and thus
the linear functions appear curved. Axis scales are identical across all
panels within column E.
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Fig. 4. EOD frequency trace of chirp `bursts' from S. terminalis. (A)
Entire burst. (B) The red-boxed portion of the burst in A is shown on an
expanded time scale. Note the plateau-like elevation of the baseline EODf and
the clustering of chirps within the burst.
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Fig. 5. EOD frequency trace of a `rasp' from Brazilian P. hasemani. (A)
Entire rasp. (B) The red-boxed portion of the rasp in A is shown on an
expanded time scale.
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Fig. 6. Discriminant function analysis (DFA) of EOD, chirp and GFR parameters.
Scatter plots of first versus second (A) and first versus
third (B) canonical roots from a DFA based on all signal parameters reveal the
ability of these roots to segregate apteronotid species. Each data point
represents a single fish, and species are color-coded as indicated. Some
species (e.g. P. hasemani and A. leptorhynchus in A, and
`A.' bonapartii and P. gimbeli in B) were
well-segregated. Other species (e.g. S. terminalis and S.
cf. roseni) overlapped considerably. (C) Performance of DFAs based on
all signal parameters, EOD parameters only, chirp parameters only, or GFR
parameters only in correctly classifying the species of individuals in
leave-one-of-each-species-out cross-validations (see Materials and methods).
The broken black line represents performance predicted based on chance alone.
All DFAs correctly classified individuals at rates exceeding chance, but DFAs
based on EODs and chirps performed better than those based on GFRs.
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Fig. 7. Correlations between signal parameters. Raw species means are shown with
the correlation calculated using TIPS model (large ; non-phylogenetic).
(A) Correlation between chirp undershoot FM and chirp–FM slope. More
negative values indicate larger undershoots (y-axis) and faster
returns to baseline EODf (x-axis). (B) Correlation between chirp FM
and chirp %AM. Larger values on the x-axis indicate greater reduction
in EOD amplitude during chirps. Letters on each data point indicate species as
follows: A, S. cf. curvirostris; B, S. cf.
roseni; C, S. terminalis; D, P. hasemani (Peru); E,
P. hasemani (Brazil); F, A. albifrons; G, A.
leptorhynchus; H, `A.' bonapartii; I, `A.' n.
sp. B; J, S. nattereri; K, S. porcinum; L, P.
gimbeli (Peru); M, P. gimbeli (Brazil); N, A.
devenanzii; O, A. balaenops.
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© The Company of Biologists Ltd 2007