Heyer, Donnelly, McDiarmid - 1994 - Measuring and monitoring biological diversity standard methods for amphibians

Prévia do material em texto

Dh·ersity 
• 
Standard Mrt hods 
ror 
fJio<llo! 4 1" 
Uoolldl._ 
~ ........ 4.u.,;IW11) 
Ill! \, \lri.OMIIil 
t ... wt .~ 
.. !h .... ,._ 
..... 
~leastl!ing 
and 
~lonitoriug; 
Biologicar 
Diversity 
• 
Standard Metlu><i 
for 
Amphibians 
Biological Divcrshy HandbOt)\.. St:rics 
Series Ecli tor: l'vlercede~ S. F'osl.ef 
1 hh series lla; been c;1ablishcu h)' the N'uti•'Ml Riologie'll Sur·•ey IU.::i. Dcpart.rn<:m of the lnlerlol'! 
on;1 1~ National Museum or ~•wrul History (Smirhsoma!l lnstlrutioo) rur the puhlit-•nQI'I o! 
mnn•wh demihn~ standatd field medlod' l'ur quahw•vc and qwm~tarive sampliag ol bmh•!Peul 
tltvor<icy. \;-4uJne:; v.>ill focus on differettt g,wps uf org;misnu.. hum plantS and artimab. l bc gu•l of 
lhe o;cnc< ,. to identify Of. where nece>s:tl')'. diwelop lh<.'<c mctl\tMb. ond proolO!e their a<h1ptam 
wurlthH.It, "'thl!l hiodiver\it.y infonnatlon will be comparable ~ern~• swdy sites. ~e.ogmpbJc arc~-., 
~nd or~nni>m~. and at the same site, ihrl)uglo Lime. 
• 
J 
Measuring 
and 
~1onitorin 
Biologica 
Diversity 
Standard ~'lethods 
for 
• • • • p- -~,Ians 
F.diwd u1 
' \\.Ronald Hrfl'l', 
Maurtl'n \. Onnnt>ll), 
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ltod \lrrrffil"' S. Fu..;tn 
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p. =· 
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Hlnloglcal tlive,.l!) ·MI>l\Uolm..,t I Hey••· 
W. Ronald 
Q,.l.M5A.M•13 1993 
5'17 (,'f\>32~'072\.-.<k)U 
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tl l ~ 'IK ~1 ot. 'iS 'l4 j ~ l 2 
tbr 7m 1\\M'WII \n ~~ i'lo.•lr~\ ~"I 
In lh., 0..01.. lllcM: .......,., ... J dut-clly •·Ill ·he: 
CiWnl'n at Utr worn. a l.l..at..1.ua L!tt 1rr:l.i\ f..tu.~ 
r-IJJ'Itkt• TbeStD•d'IV'MtUIL J""'t;.lDhrc.'t Prwi.A~\1-.N 
..,._ I'Ctll'>llra.UI>D ol[hb (,y tb.."\C oLXL.lDrl ...... 
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~h11W11nl fN Pt"nlli.I~I'IC'P or Plpc·r fnt Pl'llll"!'l.i 
lrbr..-y Maccrrol>r 7.JOA8 ·I ~81 
Tablt! xi 
Authors and Contributors xill 
Foreword xv 
Da1•id Fl, Wake. 
Prefnce xvii 
Antecedents •vii 
.'lulborship Jtvi.ii 
AcLnowledgment~ xvii1 
Cbaptu 1. lntroduction 
Previom Wort. 'Z 
lntendod Audience 3 
Contents 
Cbaptu 2. Amphibian Div~rRILy and 
I' at ural B i!.1ory: An 0\~nie!l' S 
Roy IV. McDiarmid 
intmduction S 
Order Gymnopbiona (CAecilian$) 7 
Order C'>!uilliln (Slllamandcr;) 1 
OrdGT Anura (l'rog•) 12 
Amphibian Lan• u 13 
Cbnpter 3 .1l:!;$cnlials of St•ndnrdiz:.tlon 
41\d Quantifiatlon 17 
inlroductioo I 7 
Sn>dy QUl!r.tion• 18 
1\(ltTlpling Con~•dcrations 18 
Scale 18 
Ramloml7.allon And Bias 19 
Replicatmn illld Assurnpllm~ 19 
Reponing Dal.8 'lO 
v 
Chapter 4. Rcseurch Design for Quantibltive 
Amphibian Studies 21 
Ut·AIIn C. Hayek 
lotro<Juction 2 I 
Project De&ign 2 I 
The Rc:>earch Question 21 
Fonnulacing !he Reseru-cb Hypothesis 24 
Validity 25 
Ficltl ObS<'TVation Md Stlltistical Design 28 
Data Accuracy 28 
Mr.asuremf'nt Srales R!ld Statistical 
Analysis 29 
R:mdomness 30 
lndepen<.lence 34 
Sample Size 35 
l~>tiug Euor~ 35 
Stati~tical veNus Sub,lalltive Signiflcruu:e 38 
Ch~ptu S. Keys m a SuNlessful Project: 
Associated Data and Planning 41 
lnlroduclion 4 I 
Climate and Fnvironmcnt, Mw1ha L. Crump 42 
Basic Weather Data 43 
Additional Environmental Data 44 
:Measuring Weather V;niables 45 
Autcnnated DataAcquisitioo, Charles 1?, 
J'etetsrJI'I a11d Miclu:el E. Dorr::a.s 47 
Data Loggers 47 
Euvitoomeotal Sensors 49 
Tbemtal Environment 53 
Rec(>rding Frog Call.< 54 
Recording Radiotelemetry Signals 55 
Data Standards. Roy !V. McDiarmid 57 
Geographic Characterization 58 
Habitat 58 
~ampliog Methodology 60 
Microhahitat De<c'ription, Robert r: Inger 60 
Characteristics of a Microhabirot 
Checklh~ !'II 
Basic D.:scripton: fi>r a :\olicrohabita! 
Check li't 63 
Field Method~ 66 
Vouch<'r Specimen•, /?()bert P. Reynoltf.v, 
Roffald /. Cmmbil!, and !?fry W. 
Mc:Diarmid 66 
Field Identifications 67 
Sample Size 68 
Specimen Data 69 
Call Voocbe!'s 70 
Sf'l(>.cth:lll of a Spedmen Rf'posicory 70 
Pt'TTTlibc, RtJy "'~ lvldJiarmid, Robert P. 
Rcynttld.<, and Ronald/. Cmmbie 72 
Chapter6. Standard 'fechniques for 
Inveotory and :\otunit.orin~: 75 
Sele<:tion (>f'Thchniques 75 
Standard Techniques 76 
I. Complete .Species lnvcnt(>ries. Norman 
J. Scorr. Jr. 7~ 
2. Visual &counter Survey~, Marthe! L. 
Crump nnd Nonna11J. Scou, Jr. 84 
3. Audio Strip Tronse<:ts. Barbara L. 
Zimmerman 92 
4. Quadrat Sampling. Robert G. Jae11er nnd 
R"hert F Jn:gtr 97 
5. Transect Sllll'lpling, Robert a . .Iaeger 
1()3 
6. Patch Sarupling, R"hert (,_.Iaeger I 07 
7. Str•ight·Line Drill Fence.' and Pitfall 
Traps, Paul Stephen r.,rn 1!19 
8. Sun•ey!o; .at Breeding Sites, Nm·man J. 
Smtt, .I r.. and Bru<'l! D. lVoo<f.vard I I 8 
9. Drift Fc'flces Encircling Breeding Sites, 
C. KM1U!Ih Dodd, Jr., und L>nvid 1!.. 
Score 125 
10. Quantitative Sampling (>f Amphibian 
Larvae, H. Bradky Sha}fl!r; Ros.< A. 
Alji.orrl, Bruce: D. Woodward, Stephen .1. 
1/idumJs, Ronald G.Aitig, und Clautlt 
Gascon 130 
Chapter 7. Suppl€meutal Approaches to 
Studying Amphibian Biodiversity 143 
lmroduction J 43 
Anificial Habirots 143 
• 
' 
Sampling with Arlificilll Pools, Claude 
Gascon 144 
Sampling with Anifici~l CoVl'<, (iary M. 
Fellers und Clurr!P.s A. Drost 1% 
Acouxtic Monitoring at l'ixed Sires, A. Stanley 
Rand and Oeorge /!..Drewry 150 
Target Organi~n•s and Habitnts 151 
.Background 151 
Research Design 152 
Special Considerations 152 
Tracking 152 
Thread Bobbius, \¥. Ronald Heyer I 53 
Radio Tracking, St•phen J. Ric harris, Ulrich 
Simch, and Jim> A. Alford !55 
'fr~cking with ~adioaclive Tags, Ray E. 
Ash1o11, Jr. 158 
Night Driving, H. Bradley Shajjicr and J. Edc 
Juurbock 163 
Target Organisms and tlabitats 163 
B~ckground 164 
Research Design I 65 
Field ).1ethods 165 
P~rxorincl and Materials 166 
Dauo Treatment and lnlcrprctation 166 
S~lal Consider~~rms 166 
Geographic !Jlformalion Systems 166 
GIS ann Remot~ Sensing Techniques, 
Lee-Ann C. Hayek and RtJY \V. 
McDiarmid 166 
Manual GIS Application for Habibt 
Specialists, J. Eric Jum·bock, 
SamuelS. Sweet, and Roy \V, 
Mcf)iarmitl 171 
Group Acti,•ities and Field Trips, Roy \V. 
McDiarmid ami Maureen A. iJonrlelly 
175 
1'~rg~1: Organisms and Habitats 175 
Backgroulld I 75 
Research Desib'fl I 76 
Pe~mnntl and ::VIaterial~ 176 
Da~< 'fn:alnlent and inlerpretation 176 
Special Considemliolls 176 
Exanlples of Group Activities 177 
Contems ,.ii 
Chapt~r II, ~:o1imadnll Population Si•e lS3 
lntJuduCiion 183 
Mark-Recapture, Maureen A. Don11efly and 
Craig Guye~· 1 K3 
Objectives 183 
Target Orga11isms ""'' H•bit•t 184 
Background 184 
Rese~orcllOesign and Field Methods 185 
Dat;~ Analysis and Interpretation 186 
Personnel and Materi31s 186 
l'opl~arion Estimators 186 
C3pturr.-Re~ight 200 
Bayesian Approach to Mark-Rec•pturo 
Data 200 
Removal S;!mplir.g, Lee-Ann C. Hayek 201 
Target Organisms and Habitats 201 
Background 201 
Research Llesign and Field Methods 202 
D3ta Analysis and Inte.~premrion 203 
Special C<lnsider<~tions 205 
Chapt~T 9. Analysis of Amphibian 
Hiodiversity Data 207 
l.ce-A>m C. Hayek 
Imroductioo 207 
Species Richness ).07 
Presence-Absence Data 207 
Mea.<ures of Associativn 209 
Specie.< Al>ondance 232 
Individual CountS and PropOitions 232 
lni.Crrclation;llipsof Standard Tes[S 235 
Model-Based Methods 236 
Advice about Tests for R x C Tables 237 
Advice about Tests for 2 x 2 Tables 239 
Summary and Recommcnda~ons 241 
Species Density ami Cominuously Distribun!d 
Data 243 
Gr.;phical Representa.tion 243 
Pe~-riptlve Statis~c• 244 
Spcdc:i Divi..'TI\ity 244 
(nvcntmics and Richness 245 
f\.tonituring, Rlclmess~ :md llvenncss 2.46 
S all VI~ I') JO\d Rccc'lm!n< -n<b1 11 ''" 2. ~ 1 
Nntc• ;tU 
1\p!"'nd••· Pmfl"rtit:S of A~lociorlou 
l.te:,u~• 2SS 
Eu.~.pit• f,y ln•atW~<e 2SS 
u~mplu for Lineanry Coudrtions 255 
RNI8~ C(mdlllous 261 
Cb:lpltr 10. Concluiou ood 
KmKT~~~t.:ncl.,tlom 271 
The Jrn))()IT.IIICe of [nvenlory Oar a 271 
F1exlblU1y Md lnsi&bl 272 
Modifi•lliOO of ThdmiqliCS 272 
CocnpanJOtb ""r.b p,~_. S•uut 273 
Eprlog~ '"l 
Apjlfntlll I: H:uullinJ Lht Ao1tphibiolm Z'IS 
Gtvv \I I cllc•• Uwrlcfl\ Dmst,<IIIJ II . 
HtHtUitl Hq• r 
Plulosoph~ 275 
l!Mdllll@ 275 
Aue,the•ie 276 
M~·o.t•·• 276 
Apptndl~ 2: Te<haiqutS ror Muk!na 
\mphlb.:in' 1.77 
\lo><r<ctt .-\ Do :•.xliv (''"'I G.qcr.J. F.r r 
J .. t. •to.• > "Jtd HI1XS ol A.lj!1'J 
Markl~f Adull Amphlbr&IJS 277 
l'ollttn nuopprng 277 
M~rkona 11nd Tauin& 278 
M01lun~ l.oarv•l Arnplnm.L'l'l 28:} 
R~oon.u<'oldebOO$ 2lll 
,\ llfl~nilh 3: Rctordln~ !frog C>1lls 2.85 
W Rmwltl Hc}'cr 
llqt, 0 p.nt ~· 21!5 
!lhlmr 1hc Re<: <~~dina 280 
Voucher· Sjl(' iuterls 287 
A««llleJ n~~ 287 
ApP"ndn -1: PnpArinll Alllpbilmuo.•as 
Sfitntlfie Spromul 289 
Rt>.Y IV f\1cJ)ItiJ muJ 
fniTwoJLIIC'In 280 
Docum,-n!ll100 ~ 
Spe<:omcn Prtot~"""l: 2"1 
rr-'<lu~ for KIUlnS 291 
P"""''"~VU 292 
f llli·UOO 211> 
I:J8• ;~ncll...IJ>...: ~ 
Spcco:1J l'rcp~~nW~>n• 2'15 
Padmg ll!'ltl Shll'f'IOS 295 
f\('ld Eq~ pcll<'tltlud StiJ:J'Ilts 2!-HJ 
API'C"dlc ~' CoO«tina Tis:soe for 
Hlochrork:ol Aonlym 299 
lnTf't'l F /.tri!I.>S u..J Ill Rulli]/d IJ.;u 
l"ruZ1111 Tau~ 1n II~ hdd 2"9 
Prora.ol fllf l'rtr!lnns ,.....,., 
Sample~ ~1111 
Sam pi~ S•:ro 300 
A ppcndb 6' VardoN 313 
~·lrmtf1111Sld s~ppi~H 303 
C<lmp<liCI ~o. 311 
Appendix 7: TJblf of Rando• 
1\•mbtrs .lll 
Lctv\1111 (' HuycL 
ln•lru"o'"" r, .. lJ<e ~I) 
Exrunplr H-l 
labk o(K:wd<JIU 'IUIYII>cro 311'> 
Ut..ralart Clletl J29 
lnd.t.' lSI 
.1\ddl'\''\,f, or Aulbur• ADd Conlribulors 361 
Ablolure oompliriS<>n. 234 
Ablolure difference. 234 
Abond,.,.. cu...,~. 117 
1\ccldetttal :m:ociathxt, 2.32. 
Accidental bja!, 33 
Accidental U~m(llfng. 31 
Ao:ldt~ etrecr oo d~ ..:u..oa.:y • .l9 ......,..,moo r.tc. ~ !14 
~..-.;., maoi"'""s. :Sr< .W C>IU..g 
3C\r\ity 
3>ldio <Uip ~ l«"'""vc.77, 
9~~3 
1U bJeedutg sjtf.$, 119. 120. 121. 
122-123 
•• fixed lite~ 15()...152 
ACT{lss·sitc ctJmp.uhon,118 
AetJvit}' p11Ucms, 'mtistlclll 
n:pn:-smbticm uf. C)U 
1\Jt ren'J)et<Uure, 44 • .SO 
A!po>tll«. 35-36.38 
Allrmlolive hypolhm, )S 
Alumintrm 01l'lhing. a: c1rift fb)ce 
maLtriitl, 126-127 
Amphibian 1Lr.Jmiquf':G /;Q(Jt, Z73 
Anemtlmt'b:T. 46 
cup,4B.52 
h.;ght fur, 52~~:1 
Ane·nheticsJ:;un::sthdi.z.ai:MJn. 270. 
for nuiio 1nw:miuc:r impl~~n. 157 
""!OOcliv,>'<lg. 219 
Animal care~ 162 
ANOVA. 115. 186 
,~,.,.., Sa r~ 
Aqu~r1c h»bit¥1 
dt-sl)ri:prirms uf. S<J 
J~rvilc- sam;>li.ng jn. 1 :JO-J .111 
lentic, 59 
lotle.S9 
1)11 1f\ea.!lurement in. 4:1 
qu~rlmr ~ampling in. 93 
ttu:nmm~a:r p~c;rJ'lC.O( th. 44 
Arbuoolopoda. s:ml(lllll§ or, 6 
Index 
qll3dni oamp!lns, IJ,~. 0\1 
Afthh.-es., of hcrpecnlug*u1 t1N.II.l72 
A~ spcci~ ri<:hooll/1, 24G 
A(tifi.,;iai habit»t!l, 143-151), 100 
i111ifitial t.:1tvenllr 14~, l~f.w)$0 
artiiiti<ll rot-.fua:;. 144-l.t6 
Assemblage. 32.; 
di\'t'f$ity eoo\1~ \1(, ~~ 
.OO.d """''finll ot, 125 
\lislnl ~ SI#Wty ,r(. 9fl 
A~ 
Dtckt:::os:aL 2.32 
"'"""'l<'lof, W!, WJ~)Z 
~s ut c•JUffl~cnlt,. 209 -224 
correl:\Mn UJll. 22.8 
data C)M~fi~UI'IIInd. 2:!4-22? 
homugmdty u(, 2;\U 
;ndepen<knoem.J, 227-2~~ 
lnv Mi~nce of, 230 
lini:IJ'il)' url 23f) 
mwUtrul/rmntm., 230 ~2l1 
IOOWC.mic: n:foll<mi!'J'O ~. ;n I 
351 
ASsociMiOt\ {t'mumut·dl 
1t\tOSUI'eJ <lf ( t'IJnlimlt'tft 
00111liou tllld e•,u!u3tion r.-r, 224 
;>rol"'n<es of. 229-231. 25;..27<' 
,.,,~. o(, 22!1-nll 
k:m>im lofryoof, 127- 22'1 
tndilo~al m-...asares.. P-&-!N. 
.16:l-2JG~ 
""-· Zll 
,\tJOCi\ltOO lnode\.237 
A!J\I•hPhOua, oC umpfing "rogram. 
1"-2~ 
r\Udlo 't:lp tr&llitc~.77. 92-t¥7 
Allr..,l ~utveys. Sl'f uJ:.·() Calling oll:livi~,>• 
ofbttt:diu~; siu:s, J19:. !2U, 121 , 
122.-12) 
,,utom.Jio.l d11.u .a:qoi'o-ltitJn 
e4uinmcnc. ~1-:.1 
w l<'U¢''1. 4,_..,, •7'"""¥ 
&nC "ilh catmg ¥'tivJI y tt:C«Tf.q 
~~.54-';,~ 
""' W1ih fld>O tr>o:ldng .sm....... 
~S7 
CM-irOf\mCIS)t.hl ~,nwr.;, 49~~ 
A'1tomMtd l':fdlo trx:king. ~5-S7. 
I~.S · l5R 
Autom111.«t rcccmfing, of fr<'g camnjl 
JtCdviry, 5<1 55, ~~5-287 
Ot 11>W SiltS. 150-152 
Auacm.u~ wc~fbe!" !IJJi(l(l, 48-49, 
161 
1\vttbble pnouU.Um. 12 
B.i .. 7~ "'odi£K:.Uoo cmm...c..or 
jl'1p<li>bORt><". 187.188- JR9 
M.lt, J I'll\ 
K»mmctcr 
e.on•Jnuoos ·tecO!d.ing., 46 
hl.ln!J.h\111.1. 45 
(l;anrmctnc 'J)rc.sun:, 44-4..~. 46 
BASIC pi'Qo"'TWm.lur popuhttinn ~i1.e 
onim~.tdon. ?.00 
Bli.)'CI)iWJ !lrlf)rosclt 
rn JM&r.phic on(ummioo 'l"""'" 
~liJ. f@ 
"'f<Oill>bllon sue~ .... 200 
lktl•'tlvnl' ddlamm, 2~ 
lkL» crmr, 3 (,. 3~ 
n, ... 19. lb. 2JR 
occiW:nuol 3~ 
inCOI'I"CQ\I()"'ii\1, 34 
ln~K ,)f, ~:n-23~ 
In qul*l.ltltl ~mrpling.99. 101 
Rl,.ryO.uo 
ddl.n!t lon of, ~('tS 
~"" nfr&."t."'Xi..'(ior,of. Str 
"-n~I-Jl.IOO.. me:M'!nes of 
"UinttmJ.t~l dii:trl\'lllrion, 23:\· 236, 241 
l:!inumlul mot1cl, ncg~1ivc, 24S 249 
Hinchemictll analysis. •jssue i."'tnpJa 
cullccrion for. 199 301 
Hl•loS'<>I c!ivcrsiJY, dclinitioo of. I 
81pltuio cornpl.o.\ life cy<:lc. ~ 
~~\ni, l3 
llclrCinr...,."' uocltiJIIl.1$3-l~j 
llodf_D!O. ... r.,.., «ifo:liftt'. ;13 
f'"n~' or. Sf..-57 
'RDn~>-m ·r,ot JPJn))lulg.137- J38 
Ot)~ ll~tm~lir~g. I ~:\. 136 
l:hv.n<~ntt,, "~ m~i'k:ing tcchoiqud. 
2g2-2sJ 
Ktt~•n& thorus.so. 81.92..5u ~JM, 
C~JJ.In,g aclivity 
dj11ctbufJun u(, 122. 
popow""' <~e<..ny or. <n 
8..-...,.ll; poOOd 
•"*",. •. n 
J""loo1p!. 13. 94 
ll«eo11n&Stte 
drif'l fence t:nclc>sure of. 77. ~~~-l:\H 
t.rvMI ~mpling "'· J5 
ll)e"llm ()(, 94 
$ufvt:y~ tlf, 11 ~125 
11tme Mtre.tm'.l:snd Ii\o'\'J'S, 12Z...J23 
·~ro l , J\9, 12il.121. 121-IZ! 
noc:n1 mil, 123 
rtiPOf'll, I 20 
~.120, 1 2l 
'h!U3l (Q(On"'fr t:J, as 
v.-;)ltt lew:i1b:llsUn'mero at,."' 
KrQOd..,pliQg.99. !00-IDI 
Bn1Lcn •ot.ic': ~1, l4.~ 
lht!mt:Hui.'l 
l)t'U!h ~llnlp]if,g or. 107-108 
All ot~hlm~'~ldtr hl'bit<at. 12 
l\utS..tll , ll~as ohl:illnp, 111 · 1\S, 
127-128 
0 umJw ck:nlt'itY,. 9U 
Bw•• lllld Clibsun in<h, 2.l-' 
(.llctl)J.w. 7 
'htbtw rmlir.lllOd by, S-6. 8 
'-"<ol. 7,15 
\&'"""my. 1 
CmUili,Q activit}'. J~ 
ltCOidliC ruunhori ng o( 
»t. t'm:c:rlifl¥\iTCt, 119. 12/J. J2.J, 
1:?2- 123 
11r fl :uuhitot, 15£~1.52 
•11cflo $Uip trnas~ uwpli:n;; a(, 71, 
y~ 
s.J bin~ 93.-9-t, 9-' 
.. -· fl'Ctlt$. 96 
"'•tvin>umenhtl (aclcm; m, ;&.J 45, 51 
or nu .. rumal nprdc'i1 \16 
dllri:l J> nuntm:cdin~ SCas<lt1, 12~ 
rt<Jonb:tgur. 41!, 51. 67, 97 
cquipmcm for. 54 57, 2S~-1.b(l 
rte<rnlb>i te<Mi~a•. 284-2~7 
· - S{l«im<ru, 70. 11fT 
NfC' ~I'Vl)'S<I._ 11" 
-riiOCJii>l. 95 
Call U.leftd ty. Jjij.l51 
C'.aJI n.tc. IS~ L<.J 
('fll Y(:J'ItUtion trtteJVal, l$2. 
Cnm,t.cll Sdcntific CRIO dora 
to~~r, 48 49, 51.54-~!o 
Canoe·b3sed £un·ey. 122-12J 
C. A l"l\ :RE comput~ pm ,h"l'ltm 
{Qc ~l.llatian ~iYz eslimllllott. 11!18. 
2ill, :lm. 2(\J 
f« •l'ft'•• tit<o9ty<>tlmo111on. II~ 
<'"'ll-~-. 1"'1 :lOU 
<'.1lChol>diiJ 
cq:JAilry ovorunqliJDgpcri<d. 1~. 
201 202.201 
Oflll\'&e. 140, 141 
C'll.l i.~~pcl"lLN1 ·eff-:m tslimlrtur-, 201, 
20J,205 
Co.odau.. Si!r> Salamaallcrs 
Cc:mer f(1r Hnvironmamal 0 11.111. 
MruUJJem«u, 272 
ChtnJ;{'·iR rat}O a.1im.atur. 203-lr)fi 
rwo <ttg<, 21!4-205 
Q:Jpm&l\•smtdifJOJtion t1WIIO)tQf'. d 
J"ii'o"rim me, Ill(>, IS?.II\9 
Cbl.oqa .... 209. '?21. ~ l2'1. 236. 
23~:!40 
~;:unftnttity corrcctioa oC. 141. 1.42, Z.S3 
pmblc:m~ In t.pptic:!.tiou ot, 2.11, 
23)- 2:).9 
ChiMCtone, 3~ .k.iJJin~ trututiflfl 
'291-1'11 
Cllm.alic~t.u . Sn Weather <ill~ C. 
Clu.o.:<l paJudr.rhn 
~uil.lnl sampli~ of~ 1IJ l 
"""dlim.riJo r:l. 1~4. 11!7. 20.\ 211-1 
aus,.r .....,.,.,., ~~ 
o~~~~l2.U7 
o.Iin~ 
"' ...... 45. 51 
furc(l'flpufm700databll*..11'>- 130 
ol pn:sen:.:c·ab-;cncc d"'"· 208 
ror tnr: cllppint. 279--282 
Cu(;rru.:k-nts., of l.\~!ociatiQcu. Suo 
Au.od atillns. mc~rures a( 
t.:.ollttWl!-
J"'nniu toe. 72--'l 
~ia.: c-uinctklo *· <i8 
VJ!Ico:lo<. ll3li>O ol. 7i! 
Cblnr p!Utcm. as natlm'l m21.rkcr, 
277-273 
ContbjnatiOtl framJOt:t ml;'thocl, I 1.>7 
Comp:uison. 2~~~.~ 
Complete species inH:ntor:y. 78 84 
Compul<'r 
dala c:nny ~C'.<:~uacy, 29 
field u~ nf. ftf,, J ~5 
Compuh:rinxl data mes 
of artifkill.l punl sunpling data. 146 
datat:ndingfor, 129-130 
Cum.pul~ proglams. See ulsr) 
CAPTURE com'-"'Her pcu~'Tml 
fur :tuCtlm~tcd wcatheJ 3tatlon 
operation, 41:'49 
furpupul~tion size esticaatiun, 188, 
201.202. 20) 
tOJ species dive.n.ity n.:timatinn, 116 
b.)t lnl{!ping grid dma analysi!l. 116 
Cunuept, '22 
Condilinnal di$1Tibution, 233 
Coostituti."·e dclinilir.on, 23 
CooStrl.lt.:l. ~2 
f.'outm11\•mup, ::l3 
C'.ontroJ vAriables, ::l3 
Convenit:ncc s:smpling, 3 J 
Convention nn International Trndc in 
l!nW..ngt!r..:(l Species of \\o'itd 
~<•nna and flora 1Cill:S). n, 
n 
C:mrt:clA•ion, stati-sticaL 2?.8 
Ctr\'trr, aniticial. 144. J46-J S(J 
CRC 1/undhoi'Jk of C,:t?tt.SIJS Me11wds 
jtJr Terre.1b'itll Verrelndlt:)· 
(D~-..·is). 2 
Ot~~i1tg-tJ1e-grudimt 1\{~r(lath. 
l(J+ liJ:i 
f.'nlf..o;~)\'cr, it~ mdioxth•e tag studit"s. 
1611 
Dtl"is·~'-u.Ddy comparis.oa. 20, '27:\ 
Ollh, li\'~fl:thilil)' of. 20. 271-272 
O:sts ~·t:tcy. 28-29 
Dara ocqui:iiliun syotems 
autorru~tod. 47-.'57 
ditta Jo@~ecs. 45-46, 4 7-f9 
em.:ironmcntaJ SMSOn. 49-54 
r., r.tdiu tratking, ~~ ~7 
t()f tcCOHiint ftog t.:ll.lls. ~55 
di.~rar,4f, 
P011::<~ble. 4ti 
Data analysis, 20?-276. St?e t'llr:f"' 
StMislical desigcl 
d;,t,. cla'lsi.ficatiun thf, 224 227 
asyuundcy. 2?5-2211 
cates,<Ocy lkftnhinns. 226-227 
utdt"Jing, 225 
umJert~iTIM' conrinu~. 22S 
illt~rrclufinnsb{ps of SlaJltlotn1 tesl.., in, 
235-241 
mc:~surr:mcnt scales for:. 29-:~0 
t)b;en•cr<:lft<t in. 95 
uf sp.."Cicc; a\luudact.ee datJ. 2~1-235 
of 'SJX'.dCS dMsity dab, 243 -244 
of spccjcs di.,.·":'r..ily dat~. 244 251 
of spccjes ri<'hn~'l d<n~t, 207 -232 
clttssifyitlg af &c ... 21'~227 
oonccptual re1ariun~hip~ (If, 227-
229 
mc:.sures of a.o;s1,.-,;i~tiun, 209-232 
oofafion and e-.·~:luulion, 224 
prestnOC><fh!ien(;C d:.IS~, 2U7-209 
llal~bases 
Of ~rtifit.:iotl pool Sltmpling data_ 146 
t!&a w~ing fur, 129-130 
u,,. Jogg,or, 4~6. 47-49 
(:I)YiTOIU\\Clltd SIMIIUt.'t usc:rJ With, 
49 54 
ftug calling ~oordiU.f, Oevi(;CS :.~«<.'(! 
with, 54 55, 56 
ntdiu JJackins devices usP.d with, 
Sli-.~7 
Dut:t o;hct;t 
(Ot aural SUf\o'C}'S, 1~1 
t.:npying of, :.7 
<l&:a coding fl)f, 57 
fur ti<:l<lusc, 9(1, ~1 
for larvall•.mphitticms, 6~ 
for rnalk ctcapcore :.1.udies, 1 X5-1S6 
fuT micmhabitll( data. 62, G3 
fur pitflltl trii{JJiing, 115, J 16 
fmquudrl\ts:t.mpling, 102.1r..S 
f()r sun:eys, 121, 17~. J8Q. 181. 182 
ftu systcm:ttlc s~.mpling sur;e~, 8'2 
tc;mpor.u·y. 66 
(bt O"ll:ns.cct Siimplitt.g. I 06 
f(o.r vjsual encountt.r st~T'Vt:}'S, 91 
Da!a standards. 5?~ 
fm gcC~grapttic c~cteri'l""tiun, ,')P. 
for habitats. 5~-71 
iif.lU~hc. 51) 
micmhshit;~r~. 60 66 
letrestri.al. SS-59 
tin s3mpUng., 81-84 
fur Ynuchcr l!JlCCimcns, 69-70 
Oat.> specific m<~rkin~. 16S. 277 
UC'<ld ~JlCCifll(\\S, hMdtfng uf, 11 S 
DedingAmphibi11:n P<tpulafions 
prugrum, 181 
D..""CJing :\mphibiaJ ¥urcWatiuns 
Wmkshr.p. 273 
~fJnitinn. st.:icnlitic. ~Z· 23 
Ol.•penl.le:t:t~ mudel, 2::17 
.D:pt.h ~ne. B2-133 
Utvdopm~m. uf wnphibiaus, 6 
Di()l:lel~Scamp1ing., 13 •· 145 
aft.:~~eciJi~ns. 7 
oflame. m. 134- );15, 137 
Direct dcvelupmenr. (, 
0jS1.1Jl(;e m:ili:<::r, 90 
J)j\''l:'rs.ity i.ndit.:C'I, 20, 117, 249-2~ I 
D~~A~Anulysb. 300,301 
Docuiniitltx; 24g 
DC'.Iffiinan<:e divec!li(~· (;UT\'C. 2'7 
nominance rlin:csity indioes, 249 
Ominup.c ~·stan. (If >ttudy site. 58 
tmft fr;nccs 
atbJ'"!'eLling~it.:s, n.125-J31') 
COJbfnKtiun of. 114 
tr.tp COO.S(1UCtiOnllllt.i t:-pccatJOU, 
112- 115 
ll~ wHh j.lilfa.ll tn.p<t. IJO ·91. 
1~0-117, 127 
Drmvning 
oftr3pj.lf'ci :tnimal~.l17 
of 'Kiud1ec ~ccimcns, 292 
Elfn.:( si1..c, 36 38 
G;~s 
<i.evet0$)mt:nt of, 6 
pn~ng as-spocimeu~. 291, 
294 295 
El~wic sht:ot;king. 1<1(1 
E.levatiun, uf srvdy sjte, 58 
Elevuti(IOIIJ grndiems. tranc:txt 
.".umpling of, 1m-1m 
I::.T..t:rgcnQ; lime, 5~:S, 
F.migrntjou. i>f mark·recl:lptun; srudy 
populations. J ~6 
1-:ncloJStlreS~mpling, 131.132. 
Bs-ns, 201 
l:!ndungC'fod spocjes 
tt;g>ll prote\.tion uf, ')1_ 73 
vib'U~t encmmttT;<;UCVeys of, 89 
VUUI,:hc:n;. fl6 
F.udangm::d Spoci¢'t Ace, 72 
l!nvirunm'-'nt31 g~djems. LranMa:.t 
S.""ltuj}Jin~ trf. 1 cn-tn7 
l!n-..·imnmcn("'l heaJCb. indkuo>r, 2 
Cn•·imnmen(a1 scct~r. 49-!i-l 
~·Ji.tability index, 249. Su al.l.tl 
Specie~ cquiC<'!bilil)' 
P.Tror, s(atis.lic..al 
in data a;x;urucy. 2\1 
ddinicion <Of. 26 
t)'pc 1' 3~36. :l'). 1.:t1 
lYil' !. 36. ~9 
FQ.a~ respon~. 1~7 
F.lt'tncte~. See Sptek~ 1..'\'Cnncss 
J:~pnn permit, 73 
Elc1n~llt'n. 'Jcjentjfic collecting .u)d, 
OIS 
l"al.ll\r.tliSI 7li-79.SfXulmSp~dtt 
Ji'JI 
f'•IJII•I ftlttubb.:e-,209 
Firld ""'•lllf.uo. 70 
>ltld otlivcuo >boot, ~1. 62. Snats<> 
Dtla!lx<r 
F'otki Wc:ncifo<Oiioo, 61-G:! 
f'icl~ ""'"'- 70. 11<9-'Yl 
fi'idlj '*'liC:N~Iun. rQC..TJrding of. 28-2Q 
Fldd ulps, 175 
O{.hetpeiC'IOgi~l.s.o6t.tif:t, 17~ . 
1~0- 182 
ritbi"J'.I'lml cxtimlttm·~ of pnpuliJdun 
size. 184, 187. 188. 196 
fi~t.r'vc .utllt:-st. 239. 2~'241~ '2.-42 
f"'t-.ijj1lna tcdmiques. 293--~ 
"'"'"""""' r!J"""-.. marlting 
........ ~ ll9. '283 
Fl>!t<t <AA<'I'T· ~ dxlm,..ln. 
9:1 
f-cmn.u.lill:hydc, JJ pr~crvativc, 
lQ1-m 
FORTRAN pcog~am. for pot)ulation 
size eslilWtti('>n, lK~ 
F~ritf t!H't:ie:s 
c.,..lli•n, 1 
~...U1 '<lmplm~ ol. 'XI 
llldiOII(:tive ~roc 161 
~nb), l3 
"""'""' -·i~rq labi<lJ, ~ 
""'"'' oorlca. 90 F~lion. 2.1<\ 
f'l"e:c~c·briiPUll1j:, 2.~3 
t-lrtt'7fRf,, ol tismc saa.ples, lQ9.300 
Frcqm~ncy enn·e. 90 
I"WJ: Ci.lhi. Se( CltllittJ.t at.."tivil)' 
fm;:huu..,, 143-144 
Pmgt, 1":.!· 13 
~101>\ljl ·~~>'1! 1!.'11 by. s 6. ~ -11, 13 
- ... !H• 'lildpnb 
lite c:,< lc, 14 
rqwt•fbt.11\l\! l'lduVU'~ 11-0 
lut'namy, 12 
au.,.l h<lllhln crad<i~ of, 15}-15~ 
vcx:W-lt~\iun!\ uf. P2. Srr al.VJ (~!ling 
~•l·my 
Pltnn<lt u~p 
Ct'Mt.Nt.;tion or. 114 
drif1 lfliVOW ustd with. 100, 111 , 112. 
I?W 
C<ofllpllie ~tioft. r:l !.bul} 
., •• 58 
CeoeupbJc infonnaiif.o S)·ste:~u 
(CIS),I66-175 
compmec-b.t900,16&-171 
mAnll"l, 171- 175 
=~'''"'ins and. 167, 171 17S 
OC~ii")Jhic sC3Ie.l'f sampling, 18-19 
Ol.ui bo.J bo<lcclct. .. ...oon~ 
olcria;'l1lt-279 
Ok...,.., ~. 246 
C.>&i~ ..r '""""""' •mdU:r. 18-19 
O•lYCJmmcnt~~t n:~ ':'J'u n!Jtl 
l'cmllt' 
IQr fWl•aaclivc m»teriab u~c. 1'39. 
161 1 Hi2 
Gr1tl1111tmpling, 31 
Gtadlenl!, transect umpling nf. 
103-107 
Or~o~pl111..lll \lisplay. uf ~deiO 
di .... tsily. ~244 
CrM! S)'~l21l, 89. 90 
~p O<lt\ mt..l75-l~ 
..,_lojlc>.l ""'i<·!}'litld ,.,.. 1711, 
l!lU-182 
Sf.IL~ IIMtl pnJVino: !o'UNC)'~ 1 77-~ ?ll 
OyJiUIO~'hi~ta. S« Cao::ilian'C 
Ht~:bll~f . .Set u/)·u Aquatic b.lbiuu; 
Mi.croh.lbitttl: Sublwhi1a1; 
TettestriaJ !~.abital 
~.h~~~s ia. 7tt ·79 
dl•urbw:~l'f.l!9.90 
lll~-·~c/,248 
scpk slz.e ot Sl 
Halo eft'«L 31-33 
lllll<l!J~ 
ordtll<lspecimeas.IIS 
<>( llv• '!>:<imcn< 27S...276 
IO~the:Ua fur, 2?fi 
t1f' ,,,a&-tee .. plun: spocimt'll~ Itt> 
or re(nOvJ.l sampli.ng l>po;irrum~. 
2Cl2 
ot' l!dpo1o:s. 141. !45 
dwrll1t; ~ing. 162-163 
oftr>pped ~"US, 1211 
H-""I'IH1J'Iing.31-31 
u...a...,,._ r ........... s~.....,.,.s. 
fl?, 123 
tk:.Jd.,rot\. ~ idendficadao.. 27M 
I L:.dltb.. wdaht·lt:ngth a'i inrlic~ur ,..,, 
I'Q 
Hclp'11 fnde•. 254 
f.lcrpc.c.t lll•gic;ll ~cictie& 
d411t .~~rth.ivc:." uJ'. 272 
llc1ol trips by. 17R, 1 RO · I S2 
Ueletoget)e:jL}. m.vjrunmc.nl!lJ. 11J 
lokll'to:teneit) inlli.:.s. 249 
II fooc:lloa. Su Sb.>OI'oo'<UldeJ. 
I riii'I~Jndex, 253, 154 
UiiiU)IiJ'ltm, 9(1 
of <~rtifidal puol porulfltiQn, JAG 
IU•tork•lmte.t. 26-V. 2~ 
Uome .caut,.oe.1 '.i9 
"""""'· 163- 164 
~-•l"""lry. ""ti<tica!, ZJQ, 
Z31.-ID 
UWI>i61y. tdMiV!; 44 
llumldily ....., ... 48. ro. 5.1. 
l:!yi~"Jlh.44 
Uypolhooli• 
alttroative, 35 
!Ofmlll~ioo ur. 2J. 24-2~ 
null. 3'-<16. 37. 3ti. :I?. 9~. 211\l 
lmpr~n pr.nnjt, 73 
l~dcnce, SIO!istioal. 34, 
'U7-.u8,131 
1,.1io ... "jjCcito. i!lli>IM'lwll .... m 
w..,.r;...lot!icts. or 'll'>'ir> 
..u...m.,, 249-:l51.1"' 
hwn<m<t>OIOOtl, V 
ln-t\lnte110Cl ·W1d'i~Oon dl'a:t. 
l7-2~ 
J.n(erR\edi.D.t~inte~l~ity l'e:m:h, SO 
tntcr-n~l \'alidity, 'l6-2g 
lniC:rV.tl scale, 30. W 
tnvA.fbn~. '2:3<l. 155 
ln111c,nl111fY. Su al.~o ~~.encc-&blt'J.tCt 
~ 
COIJII>Ictc tpCCies, 7&-84 
ctclinb>OO ilf, I~ 
' """11101)' d<u, ~ d, 2!1, 
l7l-VJ 
IUCN,:OSC O.XIilliqg Ampt,;b!., 
Pt~pulJitioosT<bk ruru:.2;1 
J~otl3.knlttng, :dll.ti'llicttl, 'l54 
J•W•U$81n&, '27S, 28:) 
Jolnt i~~Jtnt:y di!trributi(m, '1:U 
Jrinl P'mlu.bi1Hy 1\i.mlbulion, J)J 
IOU.YI\Cll! prugrum. JOt l"'''"blon 
<iLc <:Stim>rloo, 188 
lOLLY progmm. fi><papala-..u 
etlimlh<!n. IU 
Jut1~ S<ocil>s1ie.,.,...ar,o( 
~"""'"" ll!i, t~a. we 
Key. ~a 
Ke.y l!fll:dc::~, 'l 
Killtng. of vuud)CT l>fKCimCrt$, 
291 -29:1 
Kill· tt•wif\1:. 11 ~ 
f .. ~M~e. ~t 11lro 'Thdpoll!'~ 
.,..ld,.JnT~yol, 1-'0, 1<1 
• 
Cl»<><totkll<l of. l; 
~IQIUI.,....I>'ol. 6 
"'!>M<Iil<; Silts. j 18. 124 
11'1111tin8 o{,1ti4, l~3 
popul)tlOil dtnlity of. liB. 119 
piOOestllt~ "11 ''l)«imen~ 29L 
2\l'f-l!.J:\ 
qwul'lhlitlve 'QmpJing of. J 30-J4l 
<ll\18. II'C!)IntMI MO lntupretatlon, 
1)11-1~0 
holrt •>t-lh<lds Cot, L>L138 
..,.u.~ ot, 11~ 
t111ppu1~ oc. IJ 8 
vuu~l c:•~nlcT SW'\Ie)'S uf..&.; 
..,..,_'lp<d...,.,IS,69 
uc.n.do:, ol nudy "'"'· 5& 
Lt.t llu« 
~au ... den•'>' .,_,u 
lhermnrnda p~t m., ~ 
wrctne:)t C'ttfO"J ~ 44. q.& 
l..c=tll1c .qu.,;c htbtl~. :W 
Lite: cyc:te.. c4 llm}'hil)iafl,~, D, '" 
l.l~cllho<ld YOIIO te<l, 2~~. 239 
L(noolti tndeK.. Stt ~terstn esti:nmur 
l.inc:oln· ~>r•cllt"> lndcx. Stt Petttseu 
L""llmttlOl 
Linwily, Jn mu»'illrt'>t~f as:wcia\ic1n, 
230 
Wnc)n.n.\.CCt :;ompling, 93, lll6 
niQh\.Jrlvin,M lt(:hniA.]ue, 27, 34. 
IO:l-106 
Loc.tltcy, oft.111dy site. 58 
t.ccoJ '!'C\idJ,.<rl~ 
r~l.,.~ of.4? 
st ~~ b:'lbOC'l ID!aonarP~ 
-1111 
LA!$"""""' lftOdCI, :t.IJI-2"9 
lA'S"'"'"'• · 7J8 
~.~.,.eM tnl><kl, 241o!-2<Q 
f..ui,,WII.lftl', u( .Ow1)' ~11:, 58 
l.Atlte ~~oqu.uic hr&hi~\, !i9 
~IJIY•f'M' esthuatur, uf pvvu~U~n 
~b·"· tiJ1- tl)8 
~1.1J\ ~&ft\t)tliC UlfOOn~ni~n 
,~·gPrK."l"act{l, Hiti. l69, 
170-171 
hiatkJn~ ldt:lualt~ucs, I 84. 18:'i, 
l77 .. 21H 
hNrilrtKo 2K'l-JK3 
mfo-,)'I«'Jt'lc; IG5, 117 
au.. .. -. '""'"~ 21'1 
CUll•"'""- lSI. W 
~ "'"R'i"'· n.,- :r:s 
.......... ,.1~2~) 
.,.cllpponw, 1M. 1~ 284 
'""""""'""~.154.lS1 
:-tomc'""'p111'". n. u. n 202 
nf-•i popo!UJJ<>••· 119. IW, 
122 
c::nc:_;.w;urc ~mplb~:f. IMd, J r1 
fi•M mclhoos 0(, 1!.\S- 1 ~6 
joiW·lOi'lJiU:IIAd. 2S3 
ufl;uv ... l411 
pc::,.•uhaitm cmil("•ri('ln and, 11:$6 
pt:vulatiM c•tlnttt-ots, 1 ti6-200 
Dalley':\ n'lth1it\~.:ttdQn tJtittutror, 
1~7 . 1&~-,.y 
Ch.·wman'' mudi n.cii ftnl r.tcittttTOC, 
1~6, 1~7. IS) 
fis~cr-fold"'nn,.tor,IK4 IK7. 
1811. 194 
Jor.y-Scbcl S""'1alUbt c"rimawr, 
187. 188.200 
hW!Iy·l'lft-. 181-1811 
Pi:tct""' t$1itlw11Jr, 1114. IH'7, 
188-IK9 
Trij)le Clill:h c.<tlm"'<>r. 184. lltl. 
188.169-191 
""""""h "'"'~~" '"· 13~-· ~6 
~neq•lll·""'rh•I>II'IJ' bl•• at, I ~S 
with vi5'att cnc:uamwr ~nil'\'~~. 9t 
M3tciUogcucJ'\"it.:lc:(lt~. 211), 11;1;-l fl~. 
1;9··Z<il. Z<iz-270 
MCIUlUCtl't.OJlt, O( JiV~ 4111t}ttlbi111'1 ,, lie\ 
Mct1hitdCk's lu4elt. 246 
Metsmqrpnos;s., oC li\r;ac, a 
Me(et'OlO'JiCai ~tUi.<ln Stl \Vcttlher 
st..u\an 
M.,... •!itt. Coll.i!>liblc ol.l 
t.fo<roiiOll.ill 
"-'" lb<o. 41. 6().66 
cbe<tJiJ< f<lr. a 1-<>6 
lan·.ae saa»Ctut~pu, 13~ 
po;on "'lll!lW>II"' ur.-1~1 
fllodoalvot)' ot. 81 
lhcnnometer pfltemcrst _,, ~ 
Yi51J31 CI)ClO'IJ)I~I ~urve~ Q r, 8 R 
Micmb•bh1d .sckctJc.-•. !\f)-'" 
Mig_ratinn. tm:ecting.. $, 120 
Model! 
ror d.lt\.11. 3111tl)'!lill. 23(·-'137 
for opeu.1ive tt:~ltf.l'tTII-tuns 
D)CMUJOIIItf'f, $3--64 
Mm~(t)Tr. See (1/SQ rrecipltatlott: R3.i.n 
.lnl~'bibi.uru.' rtSf)fl(I"C 01,' 
Mobitotiu;: 
deCtlli1ilm .r. 18 
voocbcr spocin"'"' Cor. o9 
l""""O•Ioslrn•:JIIlon.l~ 
M-RC. S<t· ~tul·•"""'""' 
~lol!tplc~ Uo!ctfct ...... ~ 
~m. ._, VOUCher tpednscn 
~u.y. '!0--i\ 
ln<kx J55 
)tonlati..'V"c. &'S n~ fbCIJM.t. .VI 
Toe clippulg 
Nc:gati\•c binQmial modef. 2-i)l 
Nrt• 
bottom.B~-1.>~ 
for l11.v~d s.mpnng. B3, 134-ll~. 
1411 
Night (iriving tC<"hnjque. 2~. 34, 
16.'1-lli(i 
Nu.-:tomal r.lldio tTJcklt\g. US. I (i(J 
Noctumal ,;umplin,g 
of~cdings.itt:s. 123 
bcadlruup•fo< 97. JZI 
R!ill<dtiving tcchn;q..:. l7. ~. 
l63·f~ 
....... .)'S. 86. 89 
:"tmiJul Ink. 29. 7fJJ. V'J 
:>comcllicsc.ole. 7:07-?0l 
:"ullc=.20 
:>ullllyjlo<hc•h. 'ZS. )~~~. Ill. 209 
of J)() diff~. "\9 
st .. >p.19 
lestli nf. ~1. 38 
Nwn~H'\Jn!ll.f&incci :;arnpUn8, 71, 
so. 81. ij2~4 
Numoricat sp.;c:jes ridlllU.~. 246 
Observed effect si:c~ 37 
Oll~ccvcr, '27 
Observer bi.:u. id t.~tuu.:tr.u. 58m'f'llnJ. 
99 
Obief\.u tfftcl 9:S 
Ooe-...-JyH.ncncc iit3l)l"'" tll 
ClmJio 1~-.,(- Summ><y, 
m-tn.m 
~iaoo:! dt.ft.Oitiun. 21, , .. 
()p<rnliooal.,. Ultlu, IN 
Opt"""" «:ruperlllort. SJ- <.1 , 1 
OrdC'll'ing. of nwuenccl CLWL 2'), 'iU 
O<Oin' I «•lc, :\<1, X!7 
OticJJUI.tlun t:tudy. 12b 
O:steoJogi~ spc:cimcn~. prt.parafiou 
of. 295 
Outliu. '2~ 
0\'c:rtampliug, f.iB 
~.cli.n.~, af ~~n~us, 195-Z94 
P;mi.\•t fllltJ:!I"'.U<ct 1Ntl~aru1C'r (1-'t11, 
S1, ~19, 26.' 
rMit\<C ~ci~ iovcow.ny, \44., l-l1, 
1<6 
"""" """f''ing. 1117 I ()II 
l'm:m ""flf'iJI!i, !77-<78.1113 
I'I:Mt p>lell, lmlrodiJtg ood. w 
~73" 
~~2.1.\ 
356 TIJ1)Bl/ 
l'rnn~ 
fonoodio•al>c nwrriah ..,.._ 1~2 
~~ "'i!J'Illlc('tJII('tlfntt- 72-73 
rorselnl~~&o 140 
Pfter9M escitttiUOI. ()( P\lp!.II.J(joo site. 
18!, 187. 1&8-189 
pH. mct<~~nt:n\ uf. 4.~ 
pft nwwr, 4G. 'l 
Ph~m;Jnf:C}' 
r)f' hlecrtln(l, ·~2 
of r~oics p~re ~nd .lCOI .. ity, 90 
l'!I(>(OotloiOhr,C.O~IO . 277478 
Piclnu\.1 indr-.:.l.11 .. ~ 
r1gmau, ilUOrt'i>.1Cbf~ u mari:ilti-
nlll<llll. 27'.1. :ll\3 
~··"' .......... ma<ht. 2n415 
PIT. !!« l'>!.s<> < lnlqJOinl ..... _.., 
P•t!.U tnp. Ul6 
~t<e whh Wh\ {i,.-"Pl;C,, ~9 I, J 00--
117, 127 
"' bn:rdfn~ ''""'· 12.~1 30 
trap CQI'ltfl'\IC•ion JM opcrauon, 
11 2- 11~ 
Pit hiog, 292 
PIOtttt:;. wUb: ... huldln;;, ~t>.Su ul.111 
linnru:lhtd.~ 
Poilu t>lllpllnc, ~~-100 
Pol)•!nofP'\Ic .Sj)eCu~:•. \'()uCh!!r 
~ecirncn <M•tllm~ nr, 68. 69 
Pond I 
..._ ... "'"'PIIIi '"' JJ2·1)l, n• 
QU>diOI I><«Jlli<l.l Ill, 9 8 
vdnl),$ 
l'l>pll&lloft, d:IW••"' ... '21. 
l\lslUial1CX1 «<t<hr><. dimo<ie 
etiNitlLJn,- 4l 
I'Oflll1••1<1n dcn<ll)' $H .r!so SP«ic:i 
'k."M''il)' 
oC collli~ ntnlol. 97 
dtlft ftUCI'!/Pl(fatl trlQ) I!Q'ltt.tlioa cf, 
111 
lsn•ol, 118, 119. n~ 
or nd~rohnbif4U, ~~ 
phtitll trillfiPin~ c~matllm 11t'. 1 1 r •• 117 
lnin~.;t N~tmt'lJin;r el!Limation uf. 
10<~1~7 
Pop.tllfiOn tJTI", C•IIIU!i:lioo lllethodl 
ot, 1~1-10.~ 
~>=.w.c •lk:-ko.cd. m 
-=gl'i.II)IJ-20:1 
...od>1'<f...,ll-cNnn, 211l. 20<1 
< ............... k~ :IO~l!IS 
.... t . .....,h~. IU-.!00 
""""'""'"'!'I .. g. lUII-20.S 
l'l<:apJIIttM, ....,.,01111111 oC. <J.I. ~~. 
48,51 
~~ 
in artif .. <al pools, 146 
pitfall •roppio~ of. t29 
Pn:paBlicm 
a! spccillU:Ill, o8. 2!1!1 -2'11 
(I{ fi$SUC S3JttpiCI, 300 
l'n:$t":lcction method!:, 
n:actt\'e/intt;r;~cth~: ol'foc:-1 u(, 
l8 
P'rc$C'.no:-a~nc:r.dala, IIJ, 201 200, 
See ah·o !iinur")' dill 
of mifocilll put~ "''"'PIIn\. I"" 
Preser\·am 
of .,.ameru, II ! 
ofri<ouc...,plcs.l\16 
~·-s.m-293 
l'oiawy <ID.Il"'>>lni!M oL 20. 
m.zn 
l'!OOobiliiY '"'"PHnt. 32 
Prn::""rlunldcfin"IQn, 23-~ 
Pn~ ec\ cksign, 11-1~ 
hYJI(llht:'.li.'.l t()nnul4{~0tl ~r, 'll. 14 2:'i 
n:~an.:h tJUC.'ilh~ ~~oncl, 'll .. '%~ 
lln1t('tpl:mning, '11-7~ 
(J(cJiulmic uncJ ~n,·irunnl«tllll dat' 
<:nltet:tilm mt:lhntfs, 42-~~ 
data l':nmd!tf(l$(If, ~7 .. 06 
fur ~>gr»phlc chlll'ftlc.ri'11lc't, !li8 
f[)T tiJt-itats. :.B.-71 
foroa111pllne, 81-84 
fur \'Ot.fCbLy ~eM. 6't-"-'l 
l'mponion. Z).l 
l>imm!al di"""'l*'"nf 1.\}-ZJO 
C<larlzo!d. :!A I 
"""roc"""'" in. :U'J 
P"''l"'il iolot. 21. l6 
ProYinccwrvry, t17 1~ 170 
Psyct\:J:OJntter. 52 
<J!ng, 01.:1, Jf. 
l'uodla,l•o."l """'PIJn& b>. I lJ - I )2 
PVC pipe: 
.. ~ eud<ISttrc uw.kn:tJ, 11~ 
.., lr.lp. 128 
l')~ncm,ctcr, ~2 
Qu.adte\, li$Tldom .IL'"IS"""C.."' Of', 33 
Q.adtal d.sil)n, &&.1!8 
Qnadrm umplil!Jl, 77.97-102 
11.!1 enc:.losltre mmj)Hnr.. I :l"S 
or1.,.....,1~.B1 
P>ld• amplin~u.I07 
,..., ... ""'!~ "'""'• I~ 
Q.rutUwi.c '"""""" Wnag ftdl..;,.., 0!1-116 
c<l>n-.c. 130·141 
tkll31~wd~lcm., 
13\l-140 
r..,IQ mcllwds ror,l34-138 
~c:m:.l'l •:bls:n fOJ, 2 l·")CJ 
pmj(:ct (1~'-i gn, 'Z l 28 
.,tl\flsdcal dcsi,.2l1, 28-39 
Qu1(jetU. 234 
Rttd,·Hmilifls.~,5.....)1, 13'3. 13~1 !:8 
whh t~ttmaltn.nstuiut-rs. 1 ~6 
wllfl iruplanl>.blelr.ltl<tnillt"· l3l, 1~7 
Rldionc:ti,oe l~. tr.ldcing wilh. 
1:12-1~. 1S3 JG3 
Kain 
c!k:<l·:tn aupbibim ~J<M.;..., l 
dJcct~ealllns~\ity-'a'ot:fl' qs 
lligbtdri>~SODjl!ingdurin,!. 16~ 
RJW>II Pf!", 41. 41' 
bl!lit>g -bod.«. 48, 52 
~alk~l!4.~7 
Ra.ud~;~m nomb:r Wll~ 3~3-l, 
:H:l-31. 
RIUld<>n sampling.19. 31J, 31,l2-l4 
or PO<Ids. 131 
1-n probs.biJ11j· uwptin!. 32 
[n quadrat s~mplillg, 98 
•;tcM.tifi.:.d, 32. Su abo STrarillcatlt.nl 
ln n'Mstcr s~mpltog. 104. lflti 
Rmfoclloo. 247-248, 253 
!Wo,234 
Rillo. 234 
Rllliu ..m, :lil 
Rcx:apcure period. !&6 
R«ordiag, off~o..D.. Su c.Jiinr 
ICIIHJo. -~ ~ 
Rcc..,r"'! dt<i<;s. Sn ulw TIP> 
=>n!cr 
d~>U\ .!(, 
f·lr1~55-51 
(tJr v.~alhc-1 &t3 rec<1R'fing. >~S 46 
Rt~I-Jnl'll 5UI"Vey. 12ft 
r(e,lif'Qo'at'ln. wci,gtn-lcn.~th, l.SU 
Rtgre:ssi·:Kithreat. sl:itislt..:;at, '27 
1-tt.lhetiv~ ab\mdanct 
t1f Mrtiti.da.l (lOOI pnpulariOtts. 1-l(i 
behavl<lral foctun. in, 175 
~mali·)TI metll·xl~ ror 
audio strip Lrame<:~. 92-97 
brcorlin,g ~SUIV~:S. JJ&, .l ti. J~.tl 
drifl fencdpitfa~ll tr!'t{\5, J01, J I J 
nisho ~tivmg. 163, 166 
JYifch umpling.. t\lS 
pi/Ill '"'f1'irl!, 117 
~ ""''~'~~ 87-8!, 101 
viJ:'Al CDCOMfU' JuJTe)"'1. 9\l 
of rtW>tl<d·lttlliu .. pe<l••· 11.! 
Rbl>tm:~ ~= 
.. 
ReiWil\'C c.ncropy. of ~poojes 
~bondancc. 250 
RcJ~tHve hwnidity, 44. St•t· nlsn 
Htt~njdity srosM 
Re:ntntol!Cn,ing. J 67, 111· l7!; 
l<emn .. 'lll &.l mvling. ~ ... l:ll. 139' 
~~ 
R.q>ta "'"- 2<9, m . lSL ~ 
Rt!M~ in~ 19. ~.l4k 
Rltowch~ pro,.., d'"'l>"· 21-'l!t 
,.,ulili1if.!t1 c;1csign, 21S ~9 
0.()$ oom•cn1inn, ~~ 
tl£1111 ACCIJC:t.~. 28-29 
cl'fc<'l •iZ<, 36 -3B 
mr:sl>llrcment ''ale,, 29 · :W 
tl!n(lomncn, :W 
n~ndon'l number ta.bl~ ~3-34 
~l».li'ot.'f\CIDI uf xa:mplc' 
~3l 
.,.,p. ,.; ... 35 
W!l!'f!Ag =<lui<. 31-ll 
u!J.tinl iedcp,.dcaa. ~· 
oa~h:tinl vcrsps sub!llntit't 
•ignif~cancc-•• ;8--39 
1c,11ng errors, 35 -3~ 
l~ ()(1\\'I:C, 36, ~8. 39 
ke~earrh qu-;,tion. 21 • 22 
ltc;lll::an:h f';'.;hniquc$, s~c al.s" 
S!.m'TpUng; name."\ nt '{lf'fl/k 
n:r:l:Wqu~y 
modir~alion of, m-m 
.....-.. .... <I(. 17-IA. !7J 
~~lld-lllbil.us..,....Lrdall"' 
·~ct.l15 
Ncuau lirnt". 56 51 
Rlvon, bre.edin~ $ile san)pli.nS aJI)nv 
122-123 
N"'t1.:-roi!llif)g. 9t•. Su aiJcJ Nlg,ht 
driving teclll~que 
Rood kills. I £...'LI64, Hi6 
F: netl-c, Jmphlbi~n toaropiiJW on. S.f'" 
N'l .~tht drivi.'t. •ccbtliq_\ull 
1t "c •ob!e.. Zl6. 237. m.lol(•. 
242-141 
>a~7.1l 
h.hiw wanrion by. 5 6 .• 
Lu~!ie.7. 15 
life cyc:l~.14 
raxr~nomy. 7 
\'i)lual encuun~.:r survey!! of, K>i 
Sntu~te 
detln•11on of, 22 
f\ .. f'II''C&C:nt:.fi\*CJ)C" of. 3{)-) 1 
S11mpt-elrin: 
dc..cermr.tatio3 of, 35 
oJft~_, si7.c corrclati<ln Ctf. J6 .. 3a 
JlO' hltbi-~t. s 1 
1"8nclom samplill~ erron Jnd. 3l 
Kllttistic:ti importl'ncc ot, 209, 2.24. 
233 
(l•r tl'IMfc: $.ciimpla, :M,II 
for \'®cb<T ~iml!n M ln.:al..-, 
Coli 69 
S.wp!iog.JI-33 
~JJ.31 
u'ith aniflci.llC~>Y«. 14.4t 146-t:r;Q 
\nth artiJicia.l PI• the, 14-4-).a(, 
bouosn-net. 137-138 
b..'~x, 135.136 
bnod, 99, 11):}-JQJ 
c.luSlt'J', ~2. 247 
~nsldecllli_,m in, t)i-2(1 
wnv~nie:occ.. 3L 
dlpoct 7, 1!1, 133, IJL..IH. t:l7 
e!Do'..i<llcy. 32 
rntlome. 131.13l.I~IJS.'lel 
c~!al<oL 1~-IY 
''"" 3! h!JiflbJt:ilft!. 3)-~ 
llneuan~t. 93.106 
ni,cl!l·drivins tcw.:hni·:,uc of, '17, 34. 
1!'3-166 
tUt\hod6JI)iO' uf. 60 
ctumbec-oon:nndnctl i9, Sil, $ I, 
8~ 
pmcdiciry of. 88 
poinl. 99-100 
_rmll>!n'lily. :<1 
fllD'b\. n. •11-101 
e~~n~I3S 
ilfi01VO<-, 19 I l l 
pttl.cllllamJtling .,. \07 
transca "i-mpling "''ti'SU1, to.S 
tiuantit~tti.\•o:. 
durins; r'ield trip,, I 7.5- 176 
ofllllv•c, 13() 141 
ct«'iltdt do1gn fur. '2l-3tl 
n.tndum, 19.30. 3l.ll-)4 
for pond ..,.piing. 13J 
fcrprollabihly """'J'IntI. :r. 
fO< qoaoltJI sm~pOJ>i. 'l6 
•Mified. 32.. ,\;--'! Dl.ta S11'11tt'IOII)OI) 
fO<tr""""" """!'linw. I~. I \ltl 
•o•IOVBl. 34. 131. 1~. 2Q0-'2Il$ 
'l'lnc, 131. JJ4. 140 
shon·k'nn. ']9...84 
$10\'C pJpe, 135 
scriptr;~.nscct, 93 ·!}>~.106-107 
StuiJygoal$~nd. 1~1~ 
•yste~mujc. 32 
~·- SiblJplitogSlJNCJ>, 7?,>10. 
g!,ll24\. S-1 
t.W.x JS7 
ti.me.-cOO'itrllLDCd, 26, 'f9-80, ~4, ~~ 
II'SR,.<I,87. HS. 89.90 
a1.1djo !UtiJ). 71, tn..J)? 
othr«ding sitel, 121. 122. 123 
pl~ccmcnt of,mnic~t.$ itt, 34. l21~ 
122- tn 
.Wp, 93-9<. 106-t()) 
\'i5ull~~llf"cp..1~ Np 
U-92. 119. l'le IZ2 
Sampli'>i bias. S.t s ... 
S>mplint poiJ1b<. du•tfuutlun QO, IY 
S.mtplir~ unit,3~ 
random assignment ol, 3~-s.a 
Sc~l~. ctuuletic:al. 29--~0 
Scat~ in~u:tlity, ~5 
Sciemjfic <'UIIc:ctlns 
J:l«mib for, 12-73 
specie::\ cJUiT-tCdoo lind. ~ 
S<lnti!Wion ""'""L lj!, 16t 
s,e,rdi.limf>tQO!tr~btcd. 11) 
~sn>ll'~. <l--43 
Scm<~ 131,1)1 II(! 
permits for.7l 
·Sc)cc:Jjtxt. U1tt-r~h c cffec.:111 ol, 11-18 
Sex rnrio, oper:.tl(l(l.il 12 .. 
ShiiJOOfl'' indelt, 22?, 24~, 2Jf) :n I, 
254 
St\aJ~ClOit·We.tverinW:k , 2~ 
ShaJlClOCl-\\~inc:r 1-ndcx.. 2.'-4 
Shfpping 
of 'px.imcns. 1:96 
af li<&l<~ lOO 
~ ..... ~.??·•· 
SJ. ....... r.orr.u IT'I'IJine oe. 111 
S~ftC:mcC' fe.,.cl., '\j-).6 
S'<iJ>o(><:a.>ee ...... 209, tl1.13~ 
Stali.o;liC:JI "ers.1n Stlb'HIJin!,i\'0. !8-39 
Slhcrnilnllc, a11 tJrandicl,& I»IOM-41, 
2&3 
Si.ugle-.arell ~urvcy1 1".?0, 121 
Sketching. of "''<t' 5pccll11ti1S, 283 
.$lope ;udke,. 2:t7 
Snollt '-"till lmt;;th. 1'4~~ Z"f(J 
lK!clny fl.l4tbcSnldya t "-•11pbib\lt1:. 
""' Rq>lil<o. m 
Sad~"""''''""'" <>1. ""· 46 
So:d tt:nsperatu~ ma.-.uranet'IS Cl. 44. 
5(} 
Sctu radiation JCI19Dr. Sl 
Sp~<tld dlltll. S.l!f. C.l~og~j)h1c 
infonuatiun aytlcm (Cil~) 
Spodct. sbtandancc 
c:lldm:uion t~hniquc• (OI 
lndi>t'i.du-aJ. C<J.IIntll &lnCi rtl'n(li)Ttil)tl,, 
2:>2-233 
ir\~iatiMsbiJI' OhUu:lddllliOllC 
fiX. :23>-W 
s,..dno.....-:e(~ 
"""'•"""'~ f1>< (coni-.!) 
"'"'""'"'lll'"r. Zl-1-21~ 
I.!<Yol,lll 
tOld 'll«"liio<!I,Olual C<>tmli 
..... ""'<lilt Ql. 24<>-<148 
S(WXlc-lll·•tc.t CUf\.~ 2. .. 5>4.<11(. 
SpcckJCOUftl, 2Al. 7<44 
lll'<io'Jobundanc• ...._hip Q{, 
2116-ZAS 
Spoctf'( ,_,.,..,,!)' 
bm.d <tmol"'~ 1\(, I ~lUI 
411• llllal,ill rochni~ues lor. Z43-2A~ 
dttWIIC>II or. ?Al 
Spoclt) dl 'lnir r - ..,.,ydl tc>t!lniqutt rot. 2A4 2SJ 
dd\rut~ otj ?&a 
._. •l.ll>. 117. 2.1\J 231 
$pede< IM""""•· ;o,u. 
~Oorn.._,<ir, ___ 
2A1 
Spotlt. Oomi'Ofte ill''tniry ~ 
:,t'l 
Spctit'l "lU•t>b•l•y. Z48 
lncJ! ... Dl, 2!1~ 
Spcclt~ ov•nne11. 248-2119. 2S:l-25~ 
"'"''f!Cdr.ll dlvcr,lty c(m'lponcnf, 244 
Spa.lc~ l!ktlrttlon. !f('iC")tUic 
LOII<CIIOi 011~, ~8 
Sp<uo"' Jcotlt._..l..,_ 57 
'" "'" (.,ld.f·1~ 
~ln._,. 
•amplrlo7&-e4 
,._ ••. 1«. l•l. 146 
~ luo,'n-?1:. 2~ 2~2.., 
~~·lA 
.amploio 11M 
~ .. yi'!,t-l 
~le~ ~tutbr:f, 245 
~JI"<i<t~hnC'I1. 79.83.&7-U 
.,. .. l~6 
•• *'*'Y"" ttchniQQt~ fof. 21.11-a.n 
cLDtlt~ n·' d:ttt11 22.;s....227 
CMllp""' .......... fll!a o(, 221-22'} 
~nlr.;; or ir110rallioo. ~ 
II\Udo<l >1!6 cvtlulllllll. 2l~ 
~...,- cbllonal}'"" 
211no9 
J•rvll. t ':\ 
t.~ntpllnumc.lhi,J, inr 
l'lecdJnJ,jiN>aied. liS. 119, 17.4 
... blclii\1"" 163.166 
"'"'" <ampll"" lOll 
r«l>ll~nl'f'lns. Ill> 
~~in,:.IOI 
"''p:tlcyiA,~·c-OIDI.,..<IIL:IU 
lAS-l-1'1. HI 
~~ 
~,.... of.""Ui)Qilk:d., II~ I Jt> 
nu-rnb:l:r "'[. 3' 
JIIO.klll!;""' .mlpplug"C, II~ 19~ 4'l<l 
_..r.., rl. Cill, ZJII!..297 
do<Offiftl....,.~~~ 
or •w ....t btv.e, 29-1-29\ 
(tdOcqv;,.mc..tand "11Jik1 lOt, 
2'J6..297 
!h•Oo- l<dmiqoo>. 2~~ H4 
t111if!J a!.,..,,.,.,,., Z91~9Z 
fMt"Meu ... !PWCI)', 't\)' 
p~IC:rllatl\~ fot, ~2-~~ 
S~IS •mphlhi•na, pttt~ "~" 
oe. vs 
SSS. Su Stw<nwJC: S~io~ 
~.......,. 
s,.,.q. .. ~wY>~mmma....-,... 
m 
~"' ... ...._ 
17-11 m 
$lood.ni~ .. 7:1-141.S.O• 
•Is• .... pli"f 
....,din~,;,.. ~li!Vt)'t, I U ll5 
"nmpicl< "ffocl<O iu•Cil<OriCC. '18-8>1 
drifc fet~&.J eJ'\ti~Unu brc:«fuu,; ;.itcll, 
77. 12S-13~ 
drift It~~'" WJdl pi\J'I:IU l.f».r".77, 
'Ill •t. ICJil..ll7, I ~7 
'""'~~ll"ID!KMJ'It' untp~n,. llG-141 
pojdl-'to.:. 1~1-1()9 
...,,... .. ....,una. n. 9"1-. un 
...... , .. oe. JS-.7~. Tl 
- ....... \111. 77,tC0-107 .,..,, ........... ......, .. ,c.. 79. 
~.119.1211. 12l 
5rat< nvm 177, 1110 
Slati»r ... -:.1 dnll't 
0.1)!1 """"'""""' )6 
d•t" U:CllrKy, ~2:9 
elkl.t da..l<>-111 
J)ln,\INM:fl'lr tala., 29-.)(1 
rtcdor •tu•,)O 
"'""""' -Uiillo<. 13-.)A 
,._..,..,....., o( umJIIo'l. 30-J I 
-j$suo.3l 
•""'Jlll"!' m<lllod~ )1-3) 
Jr•t.Lrt.~ i.tt!"'r:tu.lmce, .l4 
f!AtlJUUI \ 'tr'SM itll:ttUJIUVC 
•U:uil1cancc. ll!- )'I 
"=•dnQ f'.mr• ad,. 3~-)4) 
.... """' .... 16. 1&,39 
$1>1J<OQI 10'1~ ~'~""<' oC. :J6. )$. )9 
S&lidttic ~-.e. '" s....e. ol ........... 'f'<O"'C"~ 711 71 
s.-. JI'IIC ~ m 
S~--~-SuDnll 
It_ .. 
SllllUlc>lion. 32. J>, 33. m-m 
doll ''"'Y'i! CO<. 139-1-10 
StttMU. 
llR'tdJ1: siluompltn! ol."'!· I:I'H ll 
l.,nl$1mpllfli ln. 1'13 
Smp ....,,... saJllll(in.:. 9l-94. 
1~1(11 
Slnldllnl ~ 24 
Slud-> .utt: 
~raphic chl:.raa-t'rimrJoa. ot.. S8 
loctln) oUt 
numbotr of visiu 10, "X, 7~ 
rMdolu ..!<Ctlao oC, 1~1 
Sul)l>lblll~ ealliog-m<. ~Inn 
ot '17 
Sli>lel-d.lf>o>sratft:!<.ICI.S 
~--
.. .....-~""'""" ... tar. 
ll7.lll! 
O>A1!l1t """"""'..c; 41 
S>~Yl~Jilia& :>:! 
S)•"'mlltk S~ Sorvey• (SS~). 
79.8~.81, 8~3.3' 
l'•llpolcs, 15 
arrill.cial poolsaJuplingut, 144-14~ 
dc\'dop~ue:m.al. )(l!gin.s ")'litem t t;1r, 1.C6 
"""'lllng 0\', 141. 145 
"""'· """PIW"C of. I 33 
""'""'""" 6t. 29<1, 29S 
qo......,..,~oc ,.~ us 
145 
.. _o!,JJ3 ,.. .. 
.. ~ OC'\ice, Z1'S-Z19 l:RJ 
lt!r ""boa<!M: rm;l:iD~:.l52-1~' · 
LS8-lcil 
lOt ,.wciJer $pccml~ 'l'!#l 
'l'lp: ft"(Qftloe! 
lllr coltiJlg ocll>'ity ....,.,.-m, 1o ~ 2S6 
in auJio ill'ip tr~l $l\Ufl.c~ V'7 
d.1u log:r-<:tll1lrollt'd. 48. " • '"· 
S6 
of-~mem..'XJ 
lllrllold dill ~1· SI.J,~ 
,_ p<'flaltmOO. '22 
Tauoo!ng, u nr..Uing t<ahn.iqw>, 
232-'283 
ltn.tpmllute 
lit"''. SO 
a1~na<td JDC:It5li10ilCN oil'. SfJ 
t·udy.53, ~SI 
dlh.lsol.•l 
- annu~(\lr . .o-t4 
.......,, ... n :14.57 
')('lil,-t4, :'iO 
\\OIItr, :10 
Tc:cu;inttteter, 46 
' IUrcKnial hEtbitM, dc~JiptiOil$ .Of. 
5M-5~ 
11'1trmi!ll cnviloum.u.t. 53~4 
llu:m:mUM~ 4&. .SO 
Th:-l'k. .f7 48, 50 
b14-beld. .tl 
'lho:nmso.pb. ~-!Wlfdr.i<. 
44 
Thermt'lh~cror, hlmd-llt'd., dJ. J(i 
lllennnmct..."'f 
•nA>:.immn-milli.tnnm. 43. '* 
plt~cem(';,t (If, 43-44 
Thn:~d btlhhin t.:chniquc, nf U~tck'ns, 
15>-155 
Tic~·ln·ll!nl..-inF, 1m 
lime-oonscnrintd simp)in,:. 2(,, 
1'l IIU,~ 
ftllb 'fhul ~Ain'C,., 1!:.1 
T.,..,..............,........,,.l.~ 
'r\.r!'ia~!l.'l'na!f'2Ph. ofmtfgilt poot 
JlOPU b.tirtn!, 14C\ 
11'1liUC !llimplt:!t 
colk:clion forbtochtmtital ft•talySi4. 
2t;IJ-3()1 
J\lf.ttiOM o(, 67 
p~ttvr.6<J.'l 00. 16(;. 
Pmm r(lllll tiU,. J 66 
TMds. cotllng a-ctivity measureme:.t 
t>l. Sl, 5.\ 
Tot c:UP!i~~~t.l84, 219-21:1,284 
r .... k!J.y>__..-M~by-rnlab. 
a.$('01~ii:coof. 191-19:'i 
T'nlck'it\fr. S.'S-S1, 1$2-1(~3 
r~dto I.Ill.d:ing. 155-1 SS 
rndiooclive mg. tech:.ique, 153. 
158-163 
n::ccnding lieviccs lor • .5:-i ;.7 
thrtl'1 hohhin tcd:niquc, I S\- 1 ~' 
TUn!leet K~mplinp;, R7, R&, SY, YO 
audio uril'. 77, 92-97 
O(bcudi<ll ,;t.,_IU.1Z!. 123 
pl!l«mcns of ~ur:s ir..,. ).1. t 21. 
Ill. IU 
unp.93-94,106-107 
1hn&m$-1n-hotnOf!t:DOOS 8~ 
tc!cltmquc; I ~~~lrt1 
Tno""P""dc<, P'"'"" in~<&~U><I !PIT). 
.17. 279, 2R~ 
Tntruovlfr)(e !tll.mptlng, 103 107 
Truppl!d :mim,1ls, rciCJ~5C of', t 17 
r ... .,. 
wtlphibbm~ • :mJidantt r;Y. 12(1 
'"' _,.,.., :<;1lllp!ing. 135- 1 :n 
( olrultl. I 1)1.] ll, lll. 1 14, llt. I )$ 
~m-u.p., 111 
fl'rb~ 131.138 
~fm.n 
pi\J'•U. ?1)..1)1. 109-117, I Z?, 1~6 
mn!c"lructirrn and IJ'{Itr~rion d. 
112-11~ 
with dtift ren.c:~s. 90-91 , t()') ... \1'7, 
l27 
,,.hcltcc.I2<J 
"nee flo)CS. !JU'Vat ~::Wl}lliUJ!! in, 
131 - 132 
~-Ill clrilt.tefu ..... jltJ<~J~, 
121'. 1.!7, lU. 130 
Thl>le CAI:bCilitmlo<. .C Jl')po1l1 ... AI 
s;~ 1116.1F7. 1811. 189-1?1 
,..,"'""n~ ctt~-in. r..r.o ~ .. m•or. 
2~20:. 
Twu-w11y v.willnce ~~oruUyd11. 12.$ 
2 X 2 o.hle!!, 2flS, 224, 23-fl 
lt1dellffidenre or. 22'1-221\ 
rests for, 239-242 
''Ill"'" or. 252 
Typo I enor. 35-~6. l?, 2.\l 
T:<l)e 2et'I'OT~ 36.39 
~-...- r:ill~Uoo~.- o(, 
n 
l:niled Stne!: 1-tm m Wndl.\t'(o 
Servloe. 72. 73 
V.lldity. 25-18 
mro•fl <o. 26-2R, 29 
Vtlriancc a.1alysi~.l2S 
Vendors, list or. 303-3 il 
Vr:mul pond. 5 
VF.'\.. Su.. Vi!loal F.nc.::K"In(~ Snt'C-}'1 
Vi,.J l!ttmutle< Sum:ys. 1~. "' 
114-\12 
o(-ng.U.., IIV, 1211.122 
VI'"'' k:attion,o(~-.1<0-!1 
\U:3lt:anlons. 12. Su nbl1 C•llirtP. 
liCdVi.ty 
~YIK'b« !p~!diTI<!n!l, S1 
altr:.m~Uvt t<l, 66-67 
from aniftciill pools. 14~ 
c>11 n:<onling or. 10. 2R7 
cvl.l.e«i<ln pmnh!l fur. 11 ... 73 
d:wa. a~JCb&ted w\ih, (19 10 
lar.-.1, 15 
froru [litfall<tapPn'l!o ll ,._1111. II 7 
""po~, .... p~~;., IJ)CCI<>. GS. 69 
~ of.l3!1-291 
"'PI>oi""Y r ... 10-11 
rood ldll• •s.l66 
~,mplc !tiu riYr, tJS~Y 
(<•r tissue s»mplce, '300 
nfvixval cnooW"IWt 3ut .. l!y¥. R9 
1,\-2ljsd)Jl1d 
fVr tacfinlTWl!lmitter ti(~IIIC'fl(. l56 
r"' -•i<li tog •ttacll•"'"" m 154 
Wtra c-vndoc:tivity. •W"m»l~ 
.,.,..,..._or.H 
W..ct·ootwut"""' ~ !V.nll•• 
BromtliltJ> 
WAtu tertlpt"c.ilW..re, mc,;a...,uTc:mMI 
lc:w;:l f.:Y. iO 
~&thC'r, ~ ounpf\.lbim Mc:dvl1)' t'JI(:l(lf. 
Sl 
V.~therdaua,41 1 42-4t• 
b~ic.43-44 
(f)r Sctt'.ioootl studi.-es. 43-43 
Ykatha statiotl 
~<8-4?,161 
lbu .... ii!lt>l~ (i.tcr.J:tltM "I, -14 
'M!WW>I,of~ t•li.Z7G 
\Vci1,1u-lengdt ~ 110 
Wind. c..ffi:ct qn C'JtiJing 1cdv1t} 
$.U'V~)'S. 95 
Wind dlte(:tion, 45 
WUld metcr, h~tnJ·hcl4. 4G 
\Vlnd speed. mea!us~mc:nt .,t, 4~. 4t .. 
52-!:3 
\\-'l-'!con..<:in. fmJol •nd l<lll.d 'UI'\'~ 
1"08'>1"'- 17.0. 121, 124-J\!i. 
H7 
Umenny, tul 
:t..em point.. 29 
Antecedents 
This bool: is the fin;t in a series thl!t n::c<Jm· 
men~s standardized methods for me~suring bi-
ological div~rsity. Th.ree conditions l~d to its 
prutlu~tion. For son1e tfme. Merc~des Foster 
recogni:ted the need fot such bo<>ks and was 
wifli£1g to ir.\·esr the lime needed to bring 01em 
l1> fruition. About the same rime, the R iologi-
cal Diversiry in Latin J\m~rica (BIOLAT) 
r~rogram in the Kational Mu:<t:um of Katural 
History, Smithsonian Insti!llti.:m, was seel:ing 
new area~ for program dcvelt>pmcnt. Fosttr 
p1esenterl her ideas to the BIOLAJ' steering 
committee (chaired by Runald H~yet ~nd con-
sisting uf Jonathan Coddington, KriMian 
Fauchald, Mercedes Fo~rer, Vidd Funk, and 
Preface 
Richard Vari), which <'llllJusia<lically provided 
rite re-sources tu iuirime the series. 1\1 the same 
lime, tlte scientific community was learning of 
an ostensible wmlc.hvidc d~c1int in some con-
spicuous amf)hibian populatioM in relali\'ely 
undisturbed habitats. Scientists who study am-
phibian population~ tccngnizcd tlt~. polenthrJJy 
critical nature of the ;ituation, and, under the 
guidance uf Oa\·id B. \Vakc • .some among lhem 
mobili<ed to address the issue. 
Two disrurbins problems were immediately 
encountered. Fi~4 much of Ole inlOnnation on 
amphilliall decline~ was anecdotal. Second. stan-
dardized method; criticill for <iocumenting pop-
ulation changes either did not exist OT were not 
genera11y kn(I'A·ll. Because nflhc pt..TCcivcd need 
for and inlt..-rcsl in :o;hmdudi~cd sMnpling mcth-
xvii 
od~ fol' •mphibians. thi.> group was sclcc~das 
th~ ~uhJecl of the fim book in U.e ~cries. 
l·rMcr •ct up a core commluee 10 plan the 
houk :and oversee tbe production nfa n1nnnscripr 
on amphibians. The c<l!llmiUlle included Matt· 
n:cn A. Donnelly, W. Ronald 1-l\:ycr. Rnben F. 
!Jl&l:r, •nd Roy W. McDianuid. "11><: comrmure 
alxl l'o•1er mer in April 1990 anti dectdi:d to 
contlltt individuals with "XJ>I'-"cnce witb the in-
ventOry rutd monitoring t'f amph.ibisns, m orr.k:.r 
tn request their input about !hi: value of the pro-
JMed book nnd about the technitt"'··• unci i$sues 
that should be cover~d. Ov~rull the response 
from the 35 Individuals throughnut the "odd 
who w~ll! contacted wa< pnsith e ond onrhosia.>-
lic. In !Xu:mbcr 1990 the c~ comrniuee a.~cd 
the IC"ppndeniS 10 iom1 wollilllg grouP> and •o 
draft mnnnscrlprs ID be inelu&:d. Copt~ of i!Je 
drnft manu!<cripL' w= circulnred prior to a 
workshop that was held 29-31 Much 1991 in 
Arlinw:on. Vuginia. The partioirWlts at the 
wnrkskop were Steph~n Com, Marlha Crvmp, 
Maureen IXlnneUy, Ltx·Ann H~yek:, Ronald 
lleyer, Robert JQ<:gcr, Roy McDl~nnld (Work-
tbop Cbair), Norman Scoll. h. ~radley Shaffer, 
Broce Woodward, and B~ Zlrnmtnnan. 
~ot>-.'11 ln&er was nnsble 10 !IIC1111. Crump, 
DonOJelly, Hayek, Heyer, ll!Cgcr, ..W McDinr-
mid continued working nn 1 oncl '2 Arril 1991 to 
~OIItplete IMk> i<lcotilicd dunng the work~;>. 
Jusr oofora rl1e worhhup, David Wnke asked 
the srnur dev-.l('!)ing the boo!:. to ~crve '" tile 
Working Gwup oo Pmlncols for ~~~ DecUnlng 
Amphtb•an l'opolntioos Tasl Foru: (OAPTf) of 
!be lntem>tinnal Uruon fN !be Constf'llli<lll of 
Nawre and Natural Rcsotlite$ (TUCN) Species 
Survi,111 Commission (SSC). Th!! J!TO"P il.,ureed. 
Tr.., workshop panicipants con•plcu:d three 
latge tMk• in a brief time: (1) Thl>y ldeorified 
lind receommended a sel uf slandord procedures 
for meo~uring amphibian biolngicul diversity by 
moniroring populalions .111il lnvontorying spe-
cies. Purll)' "" a resnlt of rhar process, they alsn 
dcvetOJlCd Ike essentials of the fi~ oullinc of 
dtc book. (2) They wrote all or ~11~~1llllal pal'ts 
of three chapter$. (3) They o~tlincd und :ISsumed 
responsibility fur the ~1sks needed In cmnplere 
!he bm1k. 
MaUTCeo DortncUy, Lee-Ann Hayek, Ron· 
aid Heyer, end Ro)' :McDlarn;id look on the 
Iespooalblliti"" of <;Oillpiling and cdtbna the 
publication Co~mllodons of a few nddtnonal 
e•perts \vere Wllght for the final manuscript. 
For p11blication (>UIJlCises, rhe ordel' of tile cdi. 
rors was u..:tcrmined randomly. Mercede• 
Fo~rer's etlilmial contribution 10 the boGII: so 
exceeded that uf ~ rraditlonal•erics cdil'Or •har 
!he first four <:dito~ added her"' a l'ifth editOI' 
to the vol urn.:. 
A dr:tit of the manuscript was completed in 
=ly December 1991. It was m~iewcu, te· 
vised, and <:ent for outside review in MAy 
1 !<92 The finn! dtdft WM sent to press ln Sep· 
tember 1992, 
Authorship 
The teJ<I oi 1llls book \vas developed by 11\3Jly 
indivit!tl:ll&. The edito..s as4d the chnits of' the 
"''Odting 8COilpS I() idennfy as Oul/rnn tho:le w!JO 
panicipau:d "'gntticomly in the writin11 oi the 
nlAnu.sllirt and ro ldendfy as <:lmrrit>mors rhoso 
who providud lnft)nnatioo. AU section~ for 
which authot"llhip Is oot assigned were wrirrcn 
either by the core eommitree, the editors, or the 
work~bup panieipruus as a group. 
Acknowledgments 
During the Jlt'I:JlKT:ilirm of rhis book the al.lihort 
and editoJS were aided by many people ·nte 
editor.< c•pecin\ly want to thank Sarah Klontz, 
wlto proviu<:tl •s~istance witlt tho initial m~ntl· 
script and llllcb'l""l\:'l the chapter biblmgraphte• 
into 11 Single docUrl!cnL und Robert B. Hole. Jr., 
wl1o lirelessly !luc:ntled 10 hun-'..reds of detAil• fur 
the edirors, including verification of all bil>lio-
graphic cimtions. Carolyn S. Hahn provided ex-
pen assistance in locating library rererence 
materials. Cassandra l:'hillips, Linda Wolfe, &rtd 
Fiona Wilkin.wn in~orpora!l:tJ numerous manu-
s<.:ript corrcclions into computl .. "T files and pro-
vided other a~sist<mce. Rosemary Sheffield 
provided expen advice on editorial matten;. 
ln addition Paul Usrach used his considerable 
tale111 to illuslrate the amphibian life <'>~'1" in 
Figure 1: Jennifer Shoemaker and Dali: Cn!\v· 
fnnl pn:par;;d •l.l!nwmlir.cd '"'·'ions of most of 
!lie figures; and Dale Crawford and Frances Irish 
provided artwork to enliven !he text Victor 
Krnntz took the phorograph for Figure 32. 
Charles R. Mann prepared the novel nmdom 
numbers !llble. 
RoberT lnger crirically read rhe entire manu-
script in its pennllimare version, and several 
anonymous reviewers commented on specific 
sections. Their comments helped us im-
measnrably in improving the final dcafl of the 
rook. 
Prefi;Jce xix 
Jolm Catr (Conservation lntemati<mal), 
Peter Cannell (Smithsonian ln~titution Pre.ss). 
Robert Unash (The Nature. Crmscrvancy), 
Adele Conover (Smirltsonian magazine), and 
Don Wilson (Biodiversity Program~. ~arional 
Museum of NatmaJ Ilistory) al!ended tlw 
wort:&fiop as observe-rs and pto\'i<.lcd useful 
input. 
Finally, the editor:~ want tn acknowledge the 
authnrs, cnntributor.s. and workshop patticipants 
for tlleir substantial efforts. Without rlleir r.nop-
eration and advice, this voltttne wo11ld nnt have 
beeu pos.~ible. To all of thew pcr.nns. we eJ.~;­
press our sincerest thanks. 
In~titutional support ;;nd financial $t1pport 
were provided by the U.S. Hsh and Wildlife 
Service, Department of the Tnterior: the Ameri-
can Museum of Natural Histcry; and the. 
Smitllsoni•n Institution. This book is comribu· 
tion nwnbcr 55 fmm the Biological Diversity in 
Latin America (BIOLAT) program. 
The Volume Editor.> 
(!:' 
Biolo~ical divr.r.<ity is the teml ttsed to describe 
rhe variety of aU living organixms on earth. It 
encnmpa.se• a1 least du-ee leveL> of biologicll 
otgani7.ation: genetic (individual), species, and 
ecooystem. Thix huok focu~es on biological di-
versity at the specie~ level, which is basic to the 
~tuc.ly or Ot'gfl.llism;. Snldie.o; of the abunWmcc 
;md rlisttihution of species lUive pro\'lded a 
wealth of basic knowledge, l<:d tu the de\oclo~ 
mem of " large body of ecological nnd evolu· 
tionary theory, and thrmed tlle underpituling of~ 
considerable hody of reserucb in the At>:n.' of 
ccolDgy, j.)'Stemalics., biogeography, and con~cr­
vauon biology. 
The continuing loss of biological dive.-ity 
lhtol1gb 1\nman ~ctiviticx will have a major im-
pact on our ability tu putl>ue science and to un-
Chapte•· l 
Introduction 
derstand the sdcntifk complexity of our sur-
roundings; it d!so will n~ct. the way~ in which 
eco.ystems continue to function. Thi.> impact 
im.TCaxcs the urgcnC}' for :;cientists to compt't.!-
hend exactly what ;, being lost ;and to communi· 
c.1te the.it perspecti\'e on the consc4ucnccs nf 
tlullloss to society. before the general quality of life 
on this planet ~uffcrs irrcvt..niblc dctcrion\tion. 
Knowledge of which specie:; occur in which 
aren~ is fun&mcntal tn an undt!rshmding u[ lhc 
jntriCCLcicx (\f hiulogical divcn\ity. Species list!\ 
of •ume ccm~picuou~ org.!lnisms (e.g., bunerflies. 
monkey•, birds, rutd rrees) a1·e available for 
many sires ill rbe more developea countries and 
for a few sites ill Jess de\·eloped r<:gion•, but 
comparable dara for most of the world'• biuta 
are la\.:king. Even \\'hen :<pecificdata on the tich-
l 
~~C.'S :md relntive •l'>und:mce.> r;:o( SjX'~ m a si1e 
we available,thqof~n •ruy greatlyint)-peruld 
hJlve beer. CCIII•-clod using a vuncty of proce-
dures, hinde11ll& lheJr comparabiluy. 
l.!r.cau.se <tm[>ltlbians src abundant and function-
HIIy 1mponnn1 in most fre!ihwa1er zlllil ILm:sllial 
halritm in uoplcal, '«lblrorical. ~nd 1cmpera~e 
n:gions. tbey fl1't' •ignificarn CC'fllpCincniS of the 
etnh's biota Few exampl~, sp .. 'Clc~ of Nonh 
American wood!~n<J ~alamnuders lJf I he genus 
Plerhodoll m~y t><:eur in densities nf ••vera! 
lhotJ-sand per he.:.tarc (Mecchtlm l'l72). Their 
ltJial biomass iu Wmc areas equalsli!AI of all the 
n:<idmt speclr.s ('( '""'" nmnun:.ls <Mnbined 
~nd is more th~111wicr thai of all spcci~s of birds 
during dte peak or nvinn breeding u(.1ivJty (Bur-
lOll nnd UkcM t975). Many •P"Ci~s of mnphibi · 
ans ttte wi<le rDII$iDg and ptllcntilllly could serve 
liS key >'JIC"'CS agnins1 which to evab1a111 broad 
geogn<phic or global ch211gcs ID the envin"'-
rrent. Other specie-s are habitat ~pecini!Ms oc· have 
re•lrictcd di,tributlons and coulcl s.ignnl to..-al JICT· 
curb:uiou. Certain phy"ologien!traiiS (cg" penne-
llble lokin) and ccnlugic:U uait.s (c~ amplex, 
bipllasic life cycle) moh-e ~bisn« f'OiellliaUy 
excellent indicatnt< or environment• I hcoldt. 
Because •mphcbum populations in cenain 
areas arc declilling sh><rply (B:vioaga 1990; 
Blnnsreio aoHl Woke 1990; Borchelt 1990; Phil -
bps 1990) ur ~re exhil!iling Mignilicoot local 
Ouwoabons cl'echm•nn ct al. 1991 ). we ~ggc.-st 
thai arbitral)' 11se uf pmcedures to a'·"'" the 
~~~rus of amphihi•n population• i• • luxury with 
wllic.h we should tli•flc""·~· Probable <rccie; de-
cline< provide ~o Impetus for monil<)ring the 
!ltlliUS <li' amphibi~~n populations Ill a vmety <ll. 
bium<.-. in many pllt!S of the wodd. ln this lxx>k 
we provide .,.eeorrunendation< aJJd jplidelines l{rr 
11\e inventory and mouitot'ing of amphibians. We 
pre<em a ser or ~tundanli•ed met.OOd.< who<.e use 
wifl cn.'IWe compAr:lbility between sites, at lea'<! 
wl!bin biomes. lllld lead tuw-.ud gr«~ter tmdcr· 
unnding of amphibiun specie. divcrM~ty in par-
ticular and biologJe•l diversity in gcncntl. 
We have as~ (I) !hat <!OJallll).)tl\'t d:rul 
..h(1111 nn assem~ of Oft,lollisms are m~ useful 
lh:111 qonlitativc infofll13tiM, and (2) !Mt <jWllita-
dvo inft'""''tiOII, such :Ill u li>l of specie•. lt'UIY be 
defive<J from quantltaLivc1holu. butnN tho reverse. 
Althwgh long-1enn pupolurion moniruritlg would 
oOcn he desirable, \Ye~ t!r.tin m:myases 
(•tly 'lingle im't'm<Ki<...., wtU be pa.sihle. Evm in 
thQl'e cases, sampling procedures•lmuld be quan-
lltativc, ~o tbar an initigl inventOI')' can he extended 
UtUl • monitoring flltlb'~'lllll should the nL'Cd and 
opporwn11y arise. An onventocy prutoc<ll "'flh 
<;IJGnlitalh'Ccha!acreroU..-. c:m Ol!so fadliWiceo<n-
pilliSO!lll ,..;j}J intti.,l S\ll&t':l< nfmonllllli.Jlg p<t11,'T:JmS 
Md invt:ntorics e!.sewhen;. 
Previous Work 
Much of the scte.ntlflc community h.s recog-
nized that usc of smndnrdU.cd methods i~ ~riucal 
to compllrisum of dnt~ gathered by dJI'fcrent 
workers at differeultJmc, and places (c.!j .. !)avis 
l'll\'2; Fellerse1 aL 19R~; Com and Bury 1990). 
Yet the redmiquc·rclated section.• llf publica· 
tinn~ pwmoting thi• view are dtlter brief or spe-
cific rc;> species tlf habhats. l'nr ex:unple, the 
CRC llandhoo/1: of Cmsu.< Method! for T(!n·es-
triul Vrnebtai~J (D'•v's t982) dcvoiiO< ouly sh 
p..!Jl'SIO med!od.< far .:ensm;ing :tmphiblans. 'Inc 
ilbsencoof 'tmd.aTd 1eclmiqucs is reflected in the 
work of many conservari<m, go\'ernmt..-nl, and 
<cientlfic group.' lhl!l tlfe beginning to ru;sess the 
statu., and health or amphibian populauol\s. For 
C.XUtnf'le, we &n: aW'•re or oogomg amplllbian 
monitoring PJ~l• in <1Teat Britain; the pr.l\' -
iuce of Ontario. DulB<h•; and Illinois. Kal\>:1.,, 
~nd Wiscon.,in in tho Uuitcd S.mtes. lJntiu1u-
nulely, the mcthnuoloslea used in llle6e monitor-
ing '!udies dil'fer. At pre•enr. no single nofotenc.: 
descrlbc.< r.:ctmiques for the inveruary ar moni-
••>rint~ ol ampbibb•IS ~nd llu: <-ircumstana:.< in 
which eGcb is 6pprupriate, Our intent Is to !ill 
ll•h void. 
Jntended i\ udience 
We have designed this bonk 10 meet the needs of 
con!lt:rvntloo organizationH. cnviromnemal con-
sult;~nl$. govMtm<Jlt a~tePcto• ... ; ldlife man~g­
er.;, and !<Cientim. We ~Jso hclic,·e thnt it will 
serve :.; a ttacllittl! and re~•rch •id in oolleg<!S 
aud tmtv=iti<:S 111 bed\ the oodeiJ,or.tdw!c rutd 
the grndull!c level!.. We h.we me1l 11> make d!f> 
book comprehen>ivc '"' Lhar Uldlvldu~L; with an 
uudergradu;lte clt;gn:c in scl~JIC~. ot· equivalent 
rral.t:.ing. c.1n moa.,urc (he. blologlcnl diver.;ity nf 
ruupbft>ian> UKing t'lll)' this volun"le, Wu also pro-
vide ref~renccs lo Jl(\1\lnMr li!er~rure to expedite 
dn111 analyses and f:octlitwc compruisons with 
lnfortTlllrloP g~thcrc<l in nthcr ~tLtdit.,. Mnny of 
rbe rechnlqlles tequirc ur rcwmmMd processing 
or mal'ldnl!llve amphibians and prepat'iJI& vnri· 
ous Lypc~ of vouchers. We ltave included in,! rue· 
1\(Jillllur ~'"~procedures as a ;:ri~ of :ppcntlicc,,, 
We """' anempffd to invol,•e u'{lC'1' trum 
tbrou!llt\lilt lhc world in the prcpamion of litis 
boot and to iJ>Cl!Hic cumpi<Y< !tom all g<x>-
gt'1lpllic ru"e1l.s. Ne •crtllclc"'·lhc •olur!\6 ~a!JJS a 
NCiw World bias bet~usa mo~l nf lhc o<mtnbu-
lm~ wort: In thot nte<l. Wa are confklcnl that Lhis 
bins will not lnfl\lence the ge~eral upplioahility 
of U1e methods discussed, which >huulu b~ Rp· 
propriate Cot• amphibians in all pan> uf lbe 
worl<L 
·-.. -. - -
' Q 
Amphibian Diversity and Natural 
History: An Overview 
fntrodncrion 
Few l'ttrel>rmes nre as dependent on cnvinm· 
mentnl molsnut ~s amphibians, ,.-ho.'<C geo· 
graphic: ranges. ecolog!ts, behavior><. llllcl life 
hl•tnrtc< gre strongly influencecll>y tl>e tlistribu· 
tum and abundance of water. usually in the farm 
of roU!. Jn faat, !he abrupt anal often synchro-
nl•~d breeding <1f many species offtogs with tl>c 
first rnlns i> a wtll-known natW'81 phcnnm.non. 
e~pecially in ateas where tainfaJI is di~1incUy 
>caN<mal Likewise, llle 3nnual breeding u~ra­
tions oC cerrnin north rtmpetate s>llamanolct un<l 
frog ~fl"'.~ to lnKiiticmally =<1 \'enw ponds 
nre t~ly llnk"d 10 me !ring iCc. ri'ing tempero. 
rures. and warm spring ...m. ... In cootrau. how 
r:l!nbll. hunlidir)•, pb3se$ of !he mooo, and • 
Roy W. McDiamlld 
J_-
multitucl<- of olher ecological factul'> mlet3C1 10 
affoct omphibian activity in tropiClll we1 foresrs 
is. r.nt wen understood. Just "1tett a """""c~r 
lltinlc$ tllal he or she knows ll1e "t)'Jlicnl" shus· 
non, somelhing Ullu.~ual "'unexpected happens. 
ln f!U: I, ln some complex assemb!uge• of truplcnl 
!rolf", the nonnal pattern may best be described 
as ~hao!lc or unpredictable. The complex and 
uftcn flllt\<ly unclerslood relation between the be· 
hnviar and ecology of amphibians nnd local 
wenther patterns make! designing a ~~mplin@ 
prowcnl fnr amphibians difficult. This compli · 
cn~d interdependency is something thnt U.SOtS oC 
tills bool need 1<\ k~ in mind. The three onkr> 
of living 9tDplnl>ians-<:aecili;un (Gymno-
flbjona), saJataandCTs (Caudata). and rrogs 
(AnurD)-<ue found in • v:mel}' of freshwa1er 
5 
"''""tic 8ml terrestrial environments throughout 
llw world (lohle 1). W1tedt~.r io shallow seeps on 
steep slope• ur buriuJ. in docayin.,~ or[!Mir. matetit\l 
at the booom of bow;, in deep lcaflittcrin tropit'"l 
evergreen rain forest or in bromcli"d" high in the 
forest c.mop:;. amphibians occur in nc:srly every 
kind of tm-estc~'ll and freshwater habit:!~ 
Most specie> have a biphasic, complex life 
C)'CI<" (Fig. 1). In re~pon~e 10 cert<Jin environ· 
mcnJ.al cues (<•.g., first rain of the season).tem~s· 
trial .. dults typic&!!)' move to suiUtble aquatic 
h~bic..ts ~>breed. Pull owing SOI'Il<' fonn of coutt-
ship, adult~ of oviparuus species dcp()Sit eggs in 
or near me water. 'fhese eggs hatch into free-
swimming lar..<~<' that are major cm.smners iu 
(.lqLJa.tic environments. After a pcrind of h'Tnwth, 
larvae undergo metamorphosis and move hack 
into the terrestrial environment where they t'=cd 
and continue to grow. When mature, they return 
to the aquatic environment to breed, thereby 
completing Tlle life cycle. 
Other species Lltldergo direct development, 
that is, they lack an independent larval st<Jge. 
'fheir eggs hatch into nonfeediog larvae or small 
copies of adulls. A few fomts are ovoviviparous 
or vh1ipmms. 
All understanding of the distributions and life 
histories of tbe species llnder study and know!· 
edge of lhch::sbitat~ in wllicll the organisms 
occur arc csscnLial to proj;ct &!sign and to sclcc· 
tiun of suitable s.m1p1ing methods. Such inf(rr .. 
malion al.w enhances the effectiveness of 
sampling and, thereby, the accurl!Cy with which 
runpbit>ian species richness and abuoda.uce can 
be estimated. In the following pages I briefly 
discuo.s pertinent as~ts of the life histories of 
Jiving amphihian~ and ~ummari7.e the basic hab~ 
itats n>ed by di1ferem. groups. 1 al>o provide M 
overview of the 3mphibion groops likely TO be 
Clloountcred on cl:lCh t.."\>ntincnt anll their res pee· 
live habitats (Table 1 ). Once the goal~ or a 
biodiversity project hav<' been dolincd (Chap· 
leT :3 ), data in the table sltoukllliu in the scle<-1.iun 
of suitable inventory tecl!aiques. 
Con~ider, for example, a study of frog diver· 
sity Jl a \(lw)and rdin fore~• site in South Amer· 
ica. Only 3 of 9pproximmely 10 families .iu the 
region h~ve aqu~lic adult•. and 2 of these fami· 
Iii'~ (l'ipidae. aud Pseudida<") includ<" very few 
species. Thus, aqll~tic sampling :~bould hav<" low 
priority if adults "ro tb,, U!rj,~l st&ge. However, if 
:m in\·cstig:!tor w·crc lo inventory both ::sdults :.md 
larvlic, then liquatic =rling would he "PPro-
priate. Nearly all families have aquatic L'lfV~e, 
and sampling for 11Nae might tum up purely 
aquatic adults M well. On The olhec h~nd, adults 
of som" spec.i~s in 6 families pC!Icnlially in the 
area are atboreal, and severdl of t.hese familie~ 
ha.ve high species diver~ity. A check of th<" lik•r-
aturc wuuld shuw th"t lidull• of nearly all spe-
cies in 1 of the families (Centrolenidae) me 
arh,lrcal, but only in ~treJmside \'egetation, and 
that aduh~ of many species <>f 2 other famili<"s 
(Hylidae and Leptodactylidae. e~peci~lly SJl'!· 
cie~ in the large genu~ Eleutlu?mdat:tylu.,) occur 
above ground, hut only in fore>!. Thus. The study 
design should stress techniques lhr sompling ar· 
boreal habitats, but it would al:«> have to ~pecify 
transect~ ~cro~); or ~lung streams afld tr~nsects in 
the rorest. 
to mosl instances. rcxcarchcn; win be beuer 
prepared to initistc studies if they firsr. read 
ab!>ut the gener~l ecology and life history or 
amphihi:ms likely oo be found in the are; aJtd 
crmsuh origin~l literature on speci.tk .groups. 
Excellent sulllllUiries of pertinent infmmotion 
c~n be found in general references (e.g., Duell· 
man "nd Trueh l9S6; Tyler 19&9) or in geo· 
grnpbically oriented publications (e.g., Mrka-
Schi<lt< 1975; Passmore ""d C"nutfler~ 1979; 
l.ambiris 1988, 1989; Madagascar-Ulommers· 
SchR\s>er and Bl•nc 1991; Eumpc-i\mold and 
'fl.urton 1978: Engehnruu\ et al. 1986; Asia-
&ny 1975; Inger and Stuebing !989: Mseda 
aml Matsui I\IH9; Australia--Cogger 1983; Tyler 
anti Davie.' 1986; North America-Stebbins 1985; 
Pfingstc:n and Down• 1989; Conrult and Collins 
1991; Middle America-Duellma>t 1970; VIlla 
1972; South 1\meri(:II-Oue!Jm~n 1978; Cei 
1980; Heyer ct al. 1990). 
Order Gymnophiona (caecilians) 
c-cc.lians are elonga~e. Umhlc<< II!Tlplubians; tile)' 
1aclc mils or ba'•e sbc!.t •lll:.• Th<>y un: p;ultl'Oj)kal 
ann ocrur in rues;r.lim.. ... -u:d ma!l in :ill pnrtS of the 
,,·orld cx"'!'tl-hd~ascar and the Alll>'tf~lopapuan 
rc!jion. Living caecilians are !JT'lUJ1Cd imo 6 fruni-
Les ancl 36 t-<ener..; ahuut J 65 species iU'e re<'.O~· 
t.m.n (Fro~t 19ll5. updated tlU'Ot~h 1992 by FnJ>~ 
ami Mc Diannicl. unpubl. dli!A). 
CneciliaasMedifflcult t~ ~wnplchccause.lbey 
are aquati~ or fo~·ri.sl "11C1 are mrely obsemrl. 
Because oi th!ir <ecn:ti>c habitZ. w-e know little 
aborn their a:olugy or life blstOI')'. :l.lale caeci-
lilln$ ha•c • me.rltull. proU11SlbJe r.~p•tlatory 
e>rgan. and rertilizatiOII preStiiiiBbfy is internal iu 
~II species. l\·lost c;,edli•n• urc VIViparous, al-
lhough a few are ovipar<>u>; 'l>mc hswe aquatic 
lorvae, w~iclllive in pouds or ~trewns. 
Tbe(e art> no widely known techniques r..,. 
!.alfl)lhng CDecib:tJJS. ((owever, meiliDds ror cap-
wring BqlJ'I(ic frogs. s:.lantanders. and !heir Jar-
'"" may be. useful (Ch~ptet 6). Aquatic >pedes 
muy he mnoved wilb a diproel frDm dee pel parts 
of fastA!owing creeks nttnld-elev&tions vr li>trna 
tn leaf mats andshhllow~ of mcanllcring lnwlancl 
so·eam;. l'ossorial •pcclc~< may 1:11' encollntered 
In soil beneaUI piles t'f 11111ing pl:ml n\aterials. in 
loos.e soil and fine: ~ta1.·c l .t.long streams, and 
beroenlh log>. lndividllat' o><:C~'"'no lly are cap-
ru~ in dircbes or Ott roaw. t.ml<, " ' the fmesr 
noo. allcr lrntcntial r:uns. 
Order Caudaw (salamanders) 
Adult !<alamand~~ arc terresrri:1 l. nquntic, fosso-
riul, nr arhoreol a111ph!biMS with fDur leg$ (no 
hind limhs in '5ireoids) 31ld ~ moderate w long 
rail. Most groups arc pf\ov•l•nt on the Holarctic 
region, bur • mlJ.jor radiation of pletbofi?.ntld ~al­
atnaoders wi1h direct devduponeoc ha& evolved 
Jn the New World 11Qpic,. Living salM\Qrork..-rs 
ure as'lii tled to Ill famme~. G I genera. •nd uhout 
3\10 ~[le.;ie,; (Frost 191l5, update-d tl.truugh 1992 
b)' Fnt" nrt:l .:l.lcDiannid. unpobl. datt~ &Ia. 
ll1JIOO<f"' have a variety of C<~Ut~Sltlp p.1(rcm• Md 
8n <'<!uolly dh•c:rse array of repcodli(Uvc mode." 
Viw•l nnd chemical signals seem t'l he more 
lmp~>ruo.nr lor .:onnship in ,,:lJammtders th~o in 
fro!!•· Mo~l salamanders have iotemnl fertili>;&· 
tion withour copulatinn, but a few l3r!(e •ttuatic 
spedeo h~ve ex ternal fertiUmtio~. F.11gs of 
aqumi~ •yx:cic' are laid singly. i~ stnn~;s, or in 
clumps in pr01d~ or sueams. S<>mctirne~ benoearh 
S&Jnc< or 1>113.:htd ro veg..'tlllion. ~ specie< 
boV\' •quanc larv..e lbai typic.J!IIy mct:~~norphosc 
:m;l 1110\'e to a terres1rial cm·uv.uncnr when: 
they f~e:l. grow, and onawrc. As adul~, in re-
sponse to perlint nl environmental cues, they re-
turn to aquatic t:nvironments, ofteu t11cir n1atal 
sites, lllld reproduce. Some species are p<;rm~­
oL'1111y toquatic, l\Jid l.!I!Valmctllffiorphosis Is in-
c(lm~liett (ollll&are paedomorphsJ. Ot.hers nre 
(o<:uhuth'O! pae<lornotpbs and OL'l:'.t.\lnn>l ly repro-
tlucc .-hole Maining cenain huval trllll'l. Mlll\y 
~ (mn.<t plethodontids) nre ICI'I\$Uhl , o.ie· 
JlO"l clumj)\ of eggs in moisl si~ in burmws,ln 
leaf litter. or bene!illt rocks and log~. Md hsve 
dtrect ctevelc.pment; in m<lny of there species the 
female pore~t attends the clutch. A few ~pccics 
of slllrutiAndn<i• an; ovoviviparous or vivi(l'.i· 
rouii. pt'Od\Jcinv live aquan c lorvao or (ully de-
veloped (mcwmnrpho,-e,i) youo~ 
Saltun:mders nre primarily 00<.1Wnal ""~ have 
J&:U\·ity pt>llm>< dw may ''art OOiuf.c:rc:bcally Jn. 
dh,dtl4!.\ nu~y he clumped or dlspet!oul "'i!hin ;; 
habitDt, :Uld spatisl dtstnl>unons ol\eu "illY w11h 
sex aud reproductive statu.,, Some ~ommon rem· 
pcratc species have a synchronous miJ.>T~tion from 
ctrrtsuiul huhiutt~ to AqUatic sires J.ilr n.:prudoJcrio>1. 
Bu:"""" oi dteir abundance llllrl p<>sitio>t in rhe 
fnud wei), in cermin habitats S!il&mant~ may be 
the m.:..t importnm venebrall! organi>m. For<"· 
8 nlAI'TU2 ' 
TablE 1. llahllal UbTtr.oliun b~ Amphibi:utS" 
:'llorll• 1\IDI.'rk•~ ')flddk Acnnh·l .S''"II• ;\ uocr\ta4 
1a.w~a11 Af $r ,., Po Af •• s, It Jlr; ,,, .. , Se ,.. •• -"' 
(..At£1UA "6 «J)l!'W!JflriilM-1, 
CtJCc!Ifuad:~r •• ' 
(i!.t ~rc .. ..,fl? ap.) v v 
H;ucn:aac 
<21"1<1 .. 36 "' l 
Rbiu>CilOd.- ~ A 
C! StoUt. ~l?pJ l E 
Soolccom\"'l"''l•tlr• 
(~ .v,..'l'\*='11 ~ IIPJ 
T,~ne .. 
~ """'"n "'I v 
\ rnmc.:pltlid« 
( l £Cllel), ·1 'l' ~ 
SAL.\MA~UI t\~ IC~ .. ...,.._..... • A ' A 
.Q~n..~:t-J~) ~ L 
t\CQ!biurr.uii•C' ,, 
(I t:cnu ~, .~ ,.,,) I. " 
Oyptol-n•nduf1nt " a~-a-l"• 
~Ddo: ~ A 
(J ~e:n. • .f ap.\ l 
llynuhi ul~~a:. 
(Jtl R,Cll('ta. 3!- ,p l 
·~ ' A ' • ' ~ " ' A A 
(~/ :ru'ft. "'" ~~ L e D II I> 0 0 D 
fJ«l'~d.lt • 
~2 ,GOOI:III• {t "f'·' L 
.Rhyacotri!Oiil1!!1!' " (J~.4""J l 
0\otwn.:..hlidto " A A " UJ~."l.,.!l'ltf L l 
*; if'l..'t\1~ ·' 
(2~1'!1l!,., "1~1 L 
f'!!'IX>S ,,,,...,, 
AllopluyNd" 
{l ~ll'!'lllll, i nf!,) l.'.' 
Actllroll'.i)!ldAO 
iJ e"11'1".& l~ -t') 
~ An (':'$"'·' •P• '· 
l<•t~ 
•• $, T< F• ,., 
'· 
D 
" 
'· 
A A A 
L • 
,4(rJQII~ 
••• ...... n 
~ 
• A 
L y 
1\ A A 
L lJ 
P• 
A 
A 
D 
"' 
.wi 
"' ~· T< 
A 
l. £ 
A 
L 
A • A 
L 
A A A 
L y 
F• 
A 
D 
" 
A 
0 ? 
"' "' 
AuU1'1,1111: 
.1• ,, fo At 
10 Ct4APn:.M 2 
• 
Thblot. (CqnriJwed) 
~.,~~ 1\'li!J.U.Amaik:A'' S...bA.~( 
Tuoo• ~. .Vt Tf ,.. .tl "' 
,., '• Fo "' ~. -~ Tt l'o Ar 
Bsacb,xor jlhhlld:li A 
C~L.lr.pJ 01 ...,. .. _ 
A ~ A A ~ ·' 
O.t ~M"•. ::.5n llf.l.} L L 01 L n I 
(~,JIInolw!lld~.: A A 
() JlCIIeJO, ~Uilp.) L ll I. e 
~ A ·' A " \ (6-<>.1-llsp.) 1. • l L tl.! L 
Di.(coatot•WfiC 
(.j I:;OOUI,, l ,'i ·'i*·~ 
Jlr~~·;lolw; 
(l,.....l1pJ 
Ho:~nilc(~fiD 
{\ """'11.1, 11. 'If).) 
R}tidl:lt A ·' .\ ' A " ·' .\ A A ,, ""_.... aoo 'P 1 ij BL L n "' H)>~.K 
(J!) StDrrA, '!27 ~(1.) 
l..eitt~l tnat.il..lac 
( J t,t'ID. J .rp. t 
L-<pO-t>liolw A ' ' A A ' 
_, 
·' ·' • (.50 ~ntJn, I'll ~ li!'-J L e (l L R 0 0 (lV I. 1)1 ,~ t'i OL 
lvlil!rol't)·l itill,. ·' A ' ' A A 
((>to Jl:""l~~;."'- lO&.ep ) 1. L L F' 01< !'/ 
M)i.UC,idJ.ldlc. 
ao r-on.,, u-+ o;p •• 
~luhuUclu• A ·' 
(JI pt'utl'tl, a!!)_,,) l 
l>k'<l)"<br 
(I &:AIL Z sp.} 
Pipicl •• ., A A 
(i l:ti\C'N, l7 Sll.) '· nL ...... om~ ... ·' u...-• <p.l L 
Rmi.Lo ·' A A A •' A 
(4~ .,.litH .. MY ·'S'·J L l 
Rbtco(UI~IIO 
(lft..-t&.. t91.rp..) 
Rh~kt.: • 
(l f;ru!J$, 2rp.) L h'll 
Rbtn.•P,I')·•o!J•- ·' ·' 0 leN!, J top.) L L 
A A 
L 
A A ,, A A 
L D " L ~ A 
l ,, 
L ~ 
A A 
l. L 
A ' " L E R ~ 
A A A 
L " L 
' A 
L 
A 
l 
A 
A A 
I. 
A A 
L 
·' A 
L~ 
I . 
" A L !'' 
'· • 
L 
A 
A 
N 
A 
" 
A A A A 
L DL DL 
A 
L h e 
A 
Dn 
" 
L 
A 
I< 
A 
Dl 
A } . 
BL 
A 
A 
' ti 
• 
I) 
A 
flJ.n 
A 
0 
}. II 
A A 
u !> 
A 
D 
A 
12 CJiAtJTJ~K 2 
Tabl•l. (COJJtinued) 
Sooslossidstc 
(l ~~r.•, ~ !lr.} 
" Oal~· lltul)r,•lly m:<:11ni11~ r.pe<:iet s.-e oc:crod. Geoenll>· .b .,pc..:ir ... ;,. :.~,xu~ oo1)' ooce and )s f'IS...<ign.«< to 1hc h<.lbil:tllhMI mul.t l)t»flr!l tb~· Jifc 
hi~ory \'\11.'\t; eArtj)bOIIS atC Sal81MM¢1l> lb;,t hll\~ rupulahO(Il\ t.Je\1.1 tiC ft('ll)tllli\'eJ)· D:'aleni~; :'1:111 rnpnl!ltiot•ll \bat tet'IIJS!)}' ll!l.der~O 
av.:tnm.orphe<m. f\ <l;,a;~h lli>,"'eifljjlll.ha.t~ UX£10 dot.s oot<XX.:l-'T on•hr. ~1m1im~:•1 jl'k!ialtd. 
1
' Faroil~· c'lesig:n.;nior'l$ :•n.J tl~ uun• l~u of SencJa and s·~des ful~w ""'"' ~ ll)t-.1) lL'C ufll.lnvd 0') I m (Frost and t-.-lc'DimU<I, un;~1.1bl dull•}. 
ample, in a notthern hardwood forest in New 
H•mpshire d\¢ eastern red·b;leked salamander 
(Pinhodun cinen>us) e1.ists in densities ar· 
proaching 2.50(1 rer llc<1arc anJ <'On verts energy 
into new tissue 31 about S,()()()jkcal/ha/}r. Its 
standing crop biom;o.<s uf "'hnut 1,650 g/ha is 
about 2.6 times the bioma.'" ur hircls in the area 
at the peak of their bree<ling s<:»S<m, anJ about 
equal to that of the sllrcws ancl mi<'C (Burton a!ld 
Likcnx 1975). In contrast, rettain tropical arbo-
real x~J,.mantlcrs are encoumered only rarely. 
when bmmeliad~ are removed from tree; and 
system~tically xcan:hcd. Unforcunately, thi~ 
"""'piing method also destroys their hahitat<. 
The diverxity of salamander life lli~tofies, 
behaviors, an~ habitat prefere.1ces necessitates 
a ilivcuity of methods for sampling popul~­
dons and estimating their ~i~c:-c. Fortunately. 
xaJamanders are reasouably well known and, 
except fur some tropi;;al xpccic<, c..xily sampled. 
Order Anura (frogs) 
Frog~ are jumping amphibians witl1 elongate 
hind limbs e11d no tails (all larval anuteJlS and 
male Ascapilus n·uei h~'·e t~ils). They arc the 
most diverse Md abulldllllt group of living am-
phibians, are L".Osmopolitan in distributio>t, Md 
occm· in essentially oll rerrestriollUld freshw>ter 
habitats. Their species diversity is highest in 
uopical wet forests. Taxonomically. living frogs 
are distributed runOJtg 25 families; curremly 
about 333 genera and 3.843 species nre recog-
nized (Ftost 1985, updated through 1992 by 
Fro~! and McDiarmid, unpubL tl•ta). l'rogx m~y 
he ~quotic, tcrn.:"'oial, fm;sorial. arhorc::tl, or 
some com hi nation thereof. S-ame arc cliumal, but 
tnos.t are noctum!)l Adulo. Hf most ~pecie:o are 
widely di>'P"tsccl in the enviro11ment e"cept at 
specific times of r.he year when they c<mgregate 
at ~qu~tic sir~x w hrcccl. 
v.acalh-:a(i(ln l~ .1::1.0 import.nut componem or tlte 
reproductive be.l>avior of most frogs. Am<1ng ter-
rcxtri•l vt:rtcbralc• (r<-rhaps all vertebrates). 
frogs have the highest diversity of reproductive 
behaviors and p~rental C3Te 'knm1.:n. BTeeding-
may be cxp1u:<i\'c (synchromm~ ..:Jvc-r ;;me or a 
few days at aquatic silc•) or pwlungccl (sprc•il 
over a few wcckx nr munth:< at aqu.atic ur tt:r .. 
rest rial xitcs) I Wells 1977]. Most species breed 
only once each year. btl! certain tropical fom1s 
--- ''.:.:"'.;.: ... __ _ 
• 
su 
8 lo<iluJts ~b:bl'l' qr :.nd (be. Styd 1':1 le<.. 
'Jndl,b ltlll" :wr.teail o( i:O;~l:. IO ~I brae. 
~Ia" 
- --' ---
1 tlltt>\111tc~ Aq = aqulll:1:; .\'1' • \C:JOinqut~ti~ '~= lwf'l'l'vri.ol: F(l= 1\}tl'!:Jr!;,l: At .. ttboru.l. 
t 1 J!,. )IJfiQfV mp ••d the~ ~; f\ • o111'-t.ks; !I -= qp.:: t • ~-..:: D-= -ei.u ~lopmcllz: f\ -= ~o:;; brn.c (Uic:oiOua 
Olahh~ or ~«:d.l,....._._. 1~9)'; Y =-kh:r"-c~ip3rool'1r "'~;JWWS ~"ao."!:B•~ ~a.t&A. lar--..e~ll•''t Ul 
wor 100dl6c:d ~ ·"''" P*ft) of-arLA11.~pM 'IO~l'ivlp.uwti Mdc.U:.V~'(T'fl~o! Ak;t ~pd ~1(J~.l._,O.OC: 
hli111 1;,11 •iilt ao £ io &auJtt•r )l;d.,•m sndic.,rc.. il ~M!'dl!'ll , .... t plore.:; ~ 111 1 he •lli")C haiitof 111 w),t-k ri'IC' lurva.: «:ev. "'' I r htlbe atbo1 e.J h.;. ri 1.m 
nod L jo the llijiJ:IIk h:rbb:W. rorc;t.-nJpJt", in:tirntr ll •pcclt~~ \'ohDIC l!j(t.$ P.lt ploccd I 'Ill lu-.c.• ""'~ltoio l&ivu!ol!!(, lll'l Mtehl.JI.&. Jnlp lt!C(,I Woi.IOJ ~l 
I'm). ('1'.' 01 L? illdic-.>~t:o~ lht Ml:dllil in whicltttM,.I(Ii wjrh. dire..~ &••ci')J~"'"'' <.» f;£'$ snd lt~\'iltl fmWIItJJUbly.a.:e Jo!~l, "'!1111(11 e~e Jtlkt"f.• 
arc repc>ned to breed t.hroogboutlh~ yc:u when-
eve.r ocmditions are favorable. Mos1 frogs h>vc 
¢X!Cmullertilizntiou, e4uaticeggs, nud feeding 
larvoccull ed 111Jpole<: s major rCO<'P,.IIIti2etioo of 
dlCI:~<lpoledruiuglnc!Am<l<J'hoshdistlnguiMI=. 
rrns• fromm~>St other amphibians. 
F~m~les of a rcw xpccies deposit egg• in 
le;~f Jlner or wcl mos~: 1be egg~ hatch into 
oonfeeding 1,-v:u: thnr undergo melllrnorpho-
sb .11 lhe nc•1 sue. Adults o r <~lher species 
carry .-crings of "U.~ wrapped :~ronnd lhtir 
hind leg~ m Lnlltspon tadpoles from J~rresuial 
'ues 10 aquatic HJI.cs on llleir beCk>. In a few 
sporiCll, femoles pincc t!dpole.< In water bcld 
•u broct~eliaO• ami then retnm to feed the lar-
va" with nonferlliC egg• that Jiloy deposit. 
Many species, especially ln lhc Neotropics, 
have dlrect dcvclop menc (uo frcc-sw•mmiug 
t:..n.,.} sr~ge). The..: •pecies dep~>...ll eggs in 
m<»•< ~erresuiru or Sri><trcal .Jt~: af1er &n ap-
pr<•priate peliod, lhc eggs hatch lot<> •mall 
copic• of Che •dulr. Some frog• brood eggs un 
thelt bncks; uthcro brooti er-gs or taclpul"" ln 
pm1cltes oo their b:lcks oc ~idC$, in vo.."lll 
puuchc:.~. or eveu in lhcir sromnch<. Some of 
thc.c broodlng S(l<CieS have typocn\ aquatic lar-
vne, but mnst exhibit diTc<:l developrncnt. AI · 
mosc ev<:ry repn>duct.ivc mode, including 
oonplacental viviparity and ovoviviparity, oc-
CIU3in.frog ... 
The high ~pedes dlvco.ity of frog~. thtir 
V!U'iety oi life ltiswry mo~es. and the wi~e 
QliMJftmCOI or habit.als they nSe p.:lSC a Substruc • 
ti>l1 chaUen~e IIJ the re;e<U"Cher in!=sted In 
~mpling these OTJlanisms. l·mtUJUirely. tbc 
•quacic breeding behavior o r mro~t speck~. the 
pervasiveness ur the aqnacic larval >!age, an~ 
LM species specificity of rcprwlualve 
vnc;,liutions •llow fur clh:cdve ~ling 
mcthrxls. 
Amphibian larvae 
Most runplnbiun lan•ae 'lCCW in aq;1atic babiWI~. 
Including moving W&ltr (me~ms aud rivc!'ll). 
so li water (p<lntls and lakes), aml phytotelulatu 
(tree hules. plrutt SAils and stems. hr..Jm<!lillds,