Urban birdsongs: higher minimum song frequency of an urban colonist persists in a common garden experiment

Abstract

Environmental changes caused by urbanization and noise pollution can have profound effects on acoustic communication. Many organisms use higher sound frequencies in urban environments with low-frequency noise, but the developmental and evolutionary mechanisms underlying these shifts are less clear. We used a common garden experiment to ask whether changes in minimum song frequency observed 30 years after a songbird colonized an urban environment are a consequence of behavioral flexibility or canalized changes that occur early in development. We captured male juvenile dark-eyed juncos ( Junco hyemalis thurberi ) from two recently diverged populations (urban and mountain) soon after they reached independence (aged 25-40 days), raised them in identical indoor aviaries, and studied their songs at an age of three years. We found that the large population difference in minimum frequency observed in the field persisted undiminished in the common garden despite the absence of noise. We also found some song sharing between the common garden and natal field populations, indicating that early song memorization before capture could contribute to the persistent song differences in adulthood. These results are the first to show that frequency shifts in urban birdsong are maintained in the absence of noise by genetic evolution and/or early life experiences.

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Urban birdsongs: higher minimum song frequency of an urban colonist persists
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in a common garden experiment
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373
!
374
Acknowledgments!
375
6+!%-.'N!*)!F&9+;!R5/!.$$&$%.',+!8&%-!,599+,%&'P!$5'P!/+,5/0&'P$!.'0!_)!_5+N$%/.2!*)!
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!
379
Funding
380
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384
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385

18
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#3%4K+130@#5%'15'4'5(7#+'04K1%4%&!EC59#%&5'2!3UU@)!58G1IJ!Q)!1DDH1X@)!
506
@D)! r+-2!<)4)!.'0!B)")!</&,+2!P=4$%17#'$0#5(%-$1I'$+43%1I1%-'45='%0#'3EII#33)E+'
507
I(+(51_4%1(5'()'4'5(7#+'#5718(5@#5%&!B-+!7M+/&,.'!b.%#/.9&$%2!3UU@)!164G@IJ!Q)!
508
A=1HA@3)!
509
@X)! 7%8+992!4)6)2!+%!.9)2!T(8@(54+.'K#0471(84+.'45='+1)#C013%(8-'%841%3'#Q01K1%'I(88#+4%#='
510
301)%3'15'8#+4%1(5'%('$($E+4%1(5'#3%4K+130@#5%'15'4'5(7#+'#5718(5@#5%&!B-+!
511
7M+/&,.'!b.%#/.9&$%2!3U1@)!184GDIJ!Q)!E1@XHE1DU)!
512
@[)! ^9.KK+N55/'2!_)2!<)!r+-2!.'0!F)!_#'%2!L(E5='%8453@1331(5'45='3(5,'=17#8,#5I#S'
513
4'I(@$4813(5'()'E8K45'45=')(8#3%'4I(E3%1I3&!?5'05/2!3UUX)!109G1IJ!Q)!DXHX[)!
514
@\)! ?./05$52!()?)2!+%!.9)2!L(5,'%-$#3.'3(5,'$#8)(8@45I#.'45='%0#'E3#'()'8#$#8%(18#3'
515
15'=48?C#-#='bE5I(3'ZJE5I('0-#@4+13]&!c+-.C&5/.9!E,595P;2!3UU\)!20G@IJ!Q)!\U1H
516
\UX)!
517
AU)! ?./05$52!()?)!.'0!")()!*+&,-./02!"48?C#-#='bE5I('3(5,S'+15?15,'(5%(,#5-'45='
518
)E5I%1(5'U1%0'4'$(%#5%14+'8(+#'15'8#$8(=EI%17#'13(+4%1(52!&'!L5(UK18=S'<5%#,84%17#'
519
K1(+(,-'45='#7(+E%1(548-'=17#831%-'15'%0#'bE5I(2!E)")!F+%%+/$5'!.'0!4)6)!7%8+992!
520
E0&%5/$)!3U1D2!d'&C+/$&%;!5R!?-&,.P5!</+$$J!?-&,.P5)!Q)!=1UH==X)!
521

24
A1)! b+8M.'2!:):)2!<)4)!r+-2!.'0!B)")!</&,+2!L(5,'74814%1(5'15'4'8#I#5%+-')(E5=#='
522
$($E+4%1(5'()'%0#'=48?C#-#='bE5I('ZJE5I('0-#@4+13]&!E%-595P;2!3UU[)!114G3IJ!Q)!
523
1D@H1X=)!
524
A3)! F5'&$-&2!:)2!L(5,'74814%1(5'15'4'$($E+4%1(5'()'/8#,(5'bE5I(3&!?5'05/2!1\D@)!66J!
525
Q)!@3=H@=D)!
526
A=)! c&5.,5#$%&,$!*+$+./,-!</5P/.M2!M47#5'D8(S'<5%#84I%17#'L(E5='P54+-313'
527
L()%U48#)!3U112!B-+!?5/'+99!h.K!5R!]/'&%-595P;J!Z%-.,.2!b+8!r5/N)!
528
A@)! c/#MM2!_)2!+%!.9)2!O#43E8#@#5%'48%#)4I%3'+#4='%(')4+3#'$(31%17#3'15'%0#'3%E=-'()'
529
K18=3(5,'15'5(13#&!:+%-50$!&'!E,595P;!.'0!EC59#%&5'2!3U1X)!8G11IJ!Q)!1D1XH1D3A)!
530
AA)!*&5$H?-+9+'2!7)7)2!()?)!h++2!.'0!()h)!<.%/&,+99&2!P'I(@$4813(5'K#%U##5'%U('
531
U4-3'%('@#43E8#'@151@E@')8#YE#5I-'45='45'#Q$#81@#5%4+'%#3%'()'7(I4+'$+43%1I1%-'
532
15'8#=CU15,#='K+4I?K18=3'15'8#3$(53#'%('5(13#&!c+-.C&5#/2!3U1D)!153G13IJ!Q)!
533
1@@AH1@X3)!
534
AD)! ?./05$52!()?)!.'0!4)6)!7%8+992!/5'4@$+1%E=#'45=')8#YE#5I-'15'K18=3(5,S'4'8#$+-'
535
%('*(++15,#8'#%'4+&!7'&M.9!c+-.C&5#/2!3U13)!84G@IJ!Q)!+1UH+1A)!
536
AX)! t599&'P+/2!^)7)2!+%!.9)2!/5'%0#'8#+4%1(5301$'K#%U##5.'45='@#43E8#@#5%'().'
537
4@$+1%E=#'45=')8#YE#5I-'15'K18=3(5,&!7'&M.9!c+-.C&5#/2!3U13)!84G@IJ!Q)!+1H+\)!
538
A[)! <505$2!4)2!P'$#8)(8@45I#'I(53%8415%'(5'%0#'#7(+E%1(5'()'%81++#='7(I4+1_4%1(53'15'4'
539
3(5,K18=')4@1+-'ZD433#81)(8@#3SH@K#81_1=4#]&!EC59#%&5'2!1\\X)!51G3IJ!Q)!A=XHAA1)!
540
A\)! ?./05$52!()?)2!6315,')8#YE#5I-'84%1(3'%('3%E=-'7(I4+'I(@@E51I4%1(5&!7'&M.9!
541
c+-.C&5#/2!3U1=)!85GDIJ!Q)!1A3\H1A=3)!
542
DU)!*!?5/+!B+.M2!MS'P'+45,E4,#'45='#5718(5@#5%')(8'3%4%13%1I4+'I(@$E%15,)!3U1[2!*!
543
`5#'0.%&5'!R5/!^%.%&$%&,.9!?5MQ#%&'PJ!k&+''.2!7#$%/&.)!
544

25
D1)! c.%+$2!"):)2!+%!.9)2!D4I?4,#'X@#AS'+15#48'@1Q#=C#))#I%3'@(=#+3'E315,'H1,#5'45='
545
LA&!45#/'.9!5R!^%.%&$%&,.9!^5R%8./+2!3U1AJ!Q)!1H@[)!
546
D3)! ?./05$52!()?)!.'0!4)6)!7%8+992!L048#='3(5,3'48#'()'+(U#8'$#8)(8@45I#'15'%0#'
547
=48?C#-#='bE5I(&!*5;.9!^5,&+%;!]Q+'!^,&+',+2!3U1D)!3GXI)!
548
D=)! `+-v/2!])2!+%!.9)2!"#'5(7('#3%4K+130@#5%'()'U1+=C%-$#'3(5,'IE+%E8#'15'%0#'_#K84'
549
)15I0&!b.%#/+2!3UU\)!459J!Q)!AD@HAD[)!
550
D@)! <505$2!4)2!^)F)!_#K+/2!.'0!c)!B.R%2!;18='3(5,S'%0#'15%#8)4I#'()'#7(+E%1(5'45='
551
@#I04513@&!7''#.9!*+C&+8!5R!E,595P;!EC59#%&5'!.'0!^;$%+M.%&,$2!3UU@)!35J!Q)!
552
AAH[X)!
553
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554
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555
DD)! :./9+/2!<)!.'0!^)!<+%+/$2!P'3#531%17#'$#81(=')(8'3(5,'4IYE131%1(5'15'%0#'3(5,'
556
3$488(U.'O#+(3$1_4'@#+(=14'C'4'I43#'()'4,#C+1@1%#='+#48515,&!E%-595P;2!1\[X)!
557
76G3IJ!Q)![\H1UU)!
558
DX)! :./9+/2!<)!.'0!^)!<+%+/$2!L#531%17#'$#81(=3')(8'3(5,'4IYE131%1(5')8(@'%4$#'
559
8#I(8=15,3'45='+17#'%E%(83'15'%0#'3U4@$'3$488(U.'O#+(3$1_4',#(8,1454&!E%-595P;2!
560
1\[[)!77G1IJ!Q)!XDH[@)!
561
D[)! c++,-+/2!:)")!.'0!E)7)!c/+'58&%L2!cE5I%1(54+'43$#I%3'()'3(5,'+#48515,'15'
562
3(5,K18=3&!B/+'0$!&'!E,595P;!q!EC59#%&5'2!3UUA)!20G=IJ!Q)!1@=H1@\)!
563
D\)! :#'0&'P+/2!<)?)2!;#0471(E8C,#5#%1I'454+-313'()'I4548-'3(5,S'15%#8C3%8415'
564
=1))#8#5I#3'15'3#53(8-'+#48515,.'45='#$1,#5#%1I'8E+#3&!7'&M.9!c+-.C&5#/2!1\\A)!
565
50J!Q)!1@\1H1A11)!
566

26
XU)! :+%$2!")()!.'0!:)^)!c/.&'02!`#5#%1I'74814%1(5'15%#84I%3'U1%0'#Q$#81#5I#'%('
567
=#%#8@15#'15%#815=171=E4+'=1))#8#5I#3'15'+#485#='3(5,&!</5,++0&'P$!5R!%-+!b.%&5'.9!
568
7,.0+M;!5R!^,&+',+$!5R!%-+!d'&%+0!^%.%+$!5R!7M+/&,.2!3U1[)!115G3IJ!Q)!@31H@3D)!
569
X1)! :#'0&'P+/2!<)?)!.'0!")?)!h.-%&2!dE45%1%4%17#'15%#,84%1(5'()',#5#%1I')4I%(83'15'%0#'
570
+#48515,'45='$8(=EI%1(5'()'I4548-'3(5,&!</5,++0&'P$!5R!%-+!*5;.9!^5,&+%;!cH
571
c&595P&,.9!^,&+',+$2!3U1@)!281G1X[1IJ!Q)![)!
572
X3)! *;.'2!:)4)!.'0!E)7)!c/+'58&%L2!V0#'8(+#'()'K(=-'31_#.'$0-+(,#5-.'45='4@K1#5%'
573
5(13#'15'%0#'#7(+E%1(5'()'K18='3(5,&!7M+/&,.'!b.%#/.9&$%2!1\[A)!126G1IJ!Q)![XH1UU)!
574
X=)! ?./05$52!()?)2!+%!.9)2!L(5,')8#YE#5I-'=(#3'5(%'8#)+#I%'=1))#8#5I#3'15'K(=-'31_#'
575
4@(5,'@4+#3'15'%U('(3I15#'3$#I1#3&!E%-595P;2!3UU[)!114G11IJ!Q)!1U[@H1U\=)!
576
X@)! 7%8+992!4)6)2!+%!.9)2!;(+=5#33'K#0471(8'45='3%8#33'$0-31(+(,-'15'4'5(7#+'E8K45'
577
#5718(5@#5%'3E,,#3%'84$1='I(88#+4%#='#7(+E%1(548-'4=4$%4%1(5&!c+-.C&5/.9!
578
E,595P;2!3U13)!23J!Q)!\DUH\D\)!
579
XA)! </&,+2!B)")2!<)4)!r+-2!.'0!c)!_.//2!D0#5(%-$1I'$+43%1I1%-'45='%0#'#7(+E%1(5'()'4'
580
3(I14++-'3#+#I%#='%841%')(++(U15,'I(+(51_4%1(5'()'4'5(7#+'#5718(5@#5%&!B-+!
581
7M+/&,.'!b.%#/.9&$%2!3UU[)!172J!Q)!^@\H^D3)!
582
!
583
!
584
!
585
!
586
!
587
!
588

27
Figure Captions!
589
590
Figure 1)!ET.MQ9+!$Q+,%/5P/.M$!5R!$-./+0!$5'P!%;Q+$!5K$+/C+0!K+%8++'!M5#'%.&'H!
591
.'0!#/K.'H,.Q%#/+0!M.9+$!&'!%-+!,5MM5'!P./0+')!^5'P$!72!?2!.'0!E!8+/+!Q/50#,+0!K;!
592
M5#'%.&'!M.9+$!.'0!$5'P$!c2!"2!.'0!`!8+/+!Q/50#,+0!K;!#/K.'!M.9+$)!B-+!%-&'!9&'+!5'!
593
+.,-!$Q+,%/5P/.M!M./N$!=!N_L!%5!-&P-9&P-%!R/+S#+',;!$-&R%$!K+%8++'!M5#'%.&'!.'0!
594
#/K.'!$5'P$)!!
595
!
596
Figure 2)!?5MQ./&$5'!5R!%-+!M&'&M#M2!Q+.N2!.'0!M.T&M#M!R/+S#+',&+$!5R!$5'P!%;Q+$!
597
Q/50#,+0!K;!M5#'%.&'H!.'0!#/K.'H,.Q%#/+0!M.9+$!/.&$+0!&'!.!,5MM5'!P./0+'!
598
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599
/+Q/+$+'%!/.'P+!5R!0.%.!8&%-&'!1)A!%&M+$!%-+!&'%+/S#./%&9+!/.'P+2!.'0!05%$!/+Q/+$+'%!0.%.!
600
Q5&'%$!+T,++0&'P!%-.%!/.'P+)!wwD'l!U)UU1)!
601
!
602
Figure 3)!?5MQ./&$5'!5R!%-+!M&'&M#M!G7I2!Q+.N!GcI2!.'0!M.T&M#M!G?I!R/+S#+',;!5R!
603
$5'P$!Q/50#,+0!K;!M5#'%.&'!.'0!#/K.'!M.9+$!&'!%-+!R&+90!W1\Y!.'0!&'!.!,5MM5'!P./0+'!
604
+'C&/5'M+'%)!c5TQ95%$!.$!&'!`&P)!3)!h+P+'0!&$!&0+'%&,.9!R5/!7H?)!wwD!l!U)UU1)!wD!l!U)UA)
605
606
607
608
609

28
Figure 1
610
!
611
!
612
!
613
!
614
!
615
!
616
!
617
!
618
!
619
!
620

29
Figure 2
621
!
622
!
623
!
624
!
625
!
626
!
627
!
628
!
629
******************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************
3.0
3.2
3.4
3.6
3.8
4.0
1
2
4
6
8
10
12
Minimum
Frequency
Peak
Frequency
Maximum
Frequency
Log10 Frequency (Hz)
Frequency (kHz)
Population
Mountain
Urban

30
Figure 3A
630
!
631
!
632
!
633
!
634
!
635
!
636
!
637
!
638
******************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************** ********************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************AAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAA
2.9
3.1
3.3
3.5
3.7
1
2
3
4
5
Mountain Urban
Population
Log10 Minimum Frequency (Hz)
Frequency (kHz)
Study
Aviary
Field

31
Figure 3B
639
!
640
!
641
!
642
!
643
!
644
!
645
!
646
!
647
****************************************************************************************************************************************************************************************************************************************************************************************************************************************
BBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBB
3.2
3.4
3.6
3.8
4.0
2
4
6
8
10
Mountain Urban
Population
Log10 Peak Frequency (Hz)
Frequency (kHz)

32
Figure 3C
648
!
649
!
650
!
651
!
652
!
653
!
654
!
655
!
656
****************************************************************************************************************************************************************************************************************************************************************************************************************************************
CCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCCC
3.4
3.6
3.8
4.0
4.2
4
6
8
10
12
Mountain Urban
Population
Log10 Maximum Frequency (Hz)
Frequency (kHz)

33
Supplementary Tables
657
658
Table S1.!*+$#9%$!5R!9&'+./!M&T+0!M50+9!.'.9;$+$!,5MQ./&'P!R/+S#+',;!,-./.,%+/&$%&,$!
659
K+%8++'!,5MM5'!P./0+'!Q5Q#9.%&5'$!M+.$#/+0!R/5M!Q58+/!$Q+,%/.2!/.%-+/!%-.'!R/5M!
660
$Q+,%/5P/.M$2!#$&'P!$5'P!%;Q+!.$!.!/.'05M!+RR+,%)!
661
^5'P!:+.$#/+M+'%!
E$%&M.%+!
%!
DHC.9#+!
b!
:&'&M#M!`/+S#+',;!
U)U[1!
@)A3[!
<0.001
D@!
:.T&M#M!`/+S#+',;!!
HU)UU1!
HU)U[=!
U)\=@!
AA!
!
662
Table S2)!*+$#9%$!5R!9&'+./!M&T+0!M50+9!.'.9;$+$!,5MQ./&'P!R/+S#+',;!,-./.,%+/&$%&,$!
663
K+%8++'!,5MM5'!P./0+'!Q5Q#9.%&5'$!M+.$#/+0!R/5M!$Q+,%/5P/.M$!#$&'P!&'0&C&0#.9!.$!
664
.!/.'05M!+RR+,%!/.%-+/!%-.'!$5'P!%;Q+)!!
665
^5'P!:+.$#/+M+'%!
E$%&M.%+!
%'
DHC.9#+!
b!
:&'&M#M!`/+S#+',;!
U)U\A!
=)U11!
0.003
DA!
:.T&M#M!`/+S#+',;!!
HU)U3A!
H1)@=X!
U)1A1!
DA!
<+.N!`/+S#+',;!
U)U13!
U)@3\!
U)DD[!
DA!
!
666
!
667

34
Table S3.!*+$#9%$!5R!9&'+./!M&T+0!M50+9!.'.9;$+$!,5MQ./&'P!R/+S#+',;!,-./.,%+/&$%&,$!
668
K+%8++'!,5MM5'!P./0+'!Q5Q#9.%&5'$!M+.$#/+0!R/5M!Q58+/!$Q+,%/.2!/.%-+/!%-.'!R/5M!
669
$Q+,%/5P/.M$2!#$&'P!&'0&C&0#.9!.$!.!/.'05M!+RR+,%!/.%-+/!%-.'!$5'P!%;Q+)!
670
^5'P!:+.$#/+M+'%!
E$%&M.%+!
%'
DHC.9#+!
b!
:&'&M#M!`/+S#+',;!
U)UX[!
3)13=!
0.034
D@!
:.T&M#M!`/+S#+',;!!
HU)U1=!
HU)X33!
U)@X1!
AA!
!
671
!
672
673
674
675
676
677
678
679
680
681
682
683
684

35
Figure S1.!?5//+9.%&5'!5R!G7I!M&'&M#M!.'0!GcI!M.T&M#M!R/+S#+',&+$!M+.$#/+0!C&$#.99;!
685
R/5M!%-+!$Q+,%/5P/.M!.'0!#$&'P!.!M&'#$!=U!0c!%-/+$-590!R/5M!%-+!Q58+/!$Q+,%/#M)!
686
!
687
!
688
!
689
!
690
!
691
!
692
!
693
!
694
!
695
!
696
AAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAA
3.0
3.2
3.4
3.6
3.0 3.2 3.4 3.6
Log10 Frequency (Hz) measured from Spectrogram
Log10 Frequency (Hz) measured from Power Spectrum
BBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBBB
3.7
3.8
3.9
4.0
3.7 3.8 3.9 4.0
Log10 Frequency (Hz) measured from Spectrogram
Log10 Frequency (Hz) measured from Power Spectrum

36
Figure S2. ?5MQ./&$5'!5R!%-+!$Q+,%/5P/.M!.'0!Q58+/!$Q+,%/#M!%+,-'&S#+$!R5/!
697
M+.$#/&'P!M&'&M#M!.'0!M.T&M#M!R/+S#+',;!5R!$5'P$!/+,5/0+0!&'!%-+!,5MM5'!P./0+'!
698
+'C&/5'M+'%)!E.,-!K5T!/+Q/+$+'%$!%-+!&'%+/S#./%&9+!/.'P+!.'0!M+0&.'2!8-&$N+/$!
699
/+Q/+$+'%!/.'P+!5R!0.%.!8&%-&'!1)A!%&M+$!%-+!&'%+/S#./%&9+!/.'P+2!.'0!05%$!/+Q/+$+'%!0.%.!
700
Q5&'%$!+T,++0&'P!%-.%!/.'P+)!wwD!l!U)UU1!
701
!
702
!
703
!
704
!
705
!
706
********************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************************
2.8
3.0
3.2
3.4
3.6
3.8
4.0
1
2
4
6
8
10
12
Minimum
Frequency
Maximum
Frequency
Log10 Frequency (Hz)
Frequency (kHz)
Protocol
Spectrogram
(Visual)
Power Spectrum
(Threshold)