Frohoff, T.G. 2000. Behavioral Indicators of Stress in Odontocetes During Interactions with Humans: A Preliminary Review and Discussion. International Whaling Commission Scientific Committee, SC/52/WW2.

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Fhff, T.G. 2000. Behaial Idica f Se i Odcee Dig Ieaci ih Hma:
A Pelimia Reie ad Dici. Ieaial Whalig Cmmii Scieific Cmmiee,
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Baa Ica  S  Oc
D Iac  Ha:
A Pa R a Dc
TONI G. FROHOFF
TeaMa Reeach, 321 High Sch Rd., NE, PMB 374, Baibidge Iad, WA 98110, USA
Contact email: fhff@i.ec.c
ABSTRACT
The ai ad ccece f ciabe cac ih fee-agig ceacea ha iceaed baia e he
a  decade. The e ad ae f hee ieaci a, b ae geea chaaceied b he aia iiiaig
 aig ce ad aied ii  ha (e.g., ie, ade, bae) ad aig aied  eeaed
ieaci. The deee f ga ffeig hee ieaci  he bic ha eed i a f f ieie ad
cecia eiai f dcee ad he d. Cee, e acc f haae  he aia,
icdig eee ijie ad ee aiie, hae bee eed. Ha ijie hae a iceaed, ad, i e cae, a
ha faai a dceed a a e f haaig a dhi. Thi ae ide a eiia eie f ieae
ha dce dcee behai eia idicaie f e aciaed ih ciabe ieaci beee fee-
agig dcee ad ha e he a ce. A eie f he i f h- ad g-e iac  he aia
ied i hee ieaci a a cdced a e a a aee f hee fac i igh f aagee ad
eeach. A a f 177 e ( beed feecie f he aca ccece) f e-eaed dcee behai
beed i he ce f ciabe ha ieaci ee ed i he eie. Of hee, 107 e ee ade f
ia aia. Eigh-ee (a 50%) f he a e ied ie . Thi high i f e-
eaed ee eed f ie ae idicae ha hei iac  dcee a be ei de a aie f
cdii. Of he 38 behai e ed, aidace a he  fee eed, fed b a ad ahe
agai he ae ad fcef hig ad biig ie. Thi eie a deae ha dce ehibiig
he highe degee f cac ih ha ae geea a he geae i f ij, ie, ad deah. I aica,
icide i hich ha ieia ijed  ied aia ied i ciabe ieaci ee eed a
ecie f ia aia ad aia ega iied ih fd. The eeach  hi bjec iae h
ideificai f e-eaed behai i aabe i deeiig iac  aia a e a iiiig iac he
icaed i aagee ga. Hee, eeach ha age bee eiced  die f h-e iac 
he aia. Becae f he difficie ihee i beig behaia iac  dcee, deeiai f  
iia iac hd  be ade baed  he abece f eceied h-e ee ae. Reeach, aagee
ad egiai f hee ga hae agged fa behid hei eai. Lg-e, aiaie die f ceacea
ageed f eceaia ha acii eed  be iiiaed a ea a ibe i he deee f he acii.
Maagee aeaed  be  effecie he i a baed  eeach, a ie- ad aia-ecific ad a
ieeed ea i he deee f he ieaci. A, adeae iig, edcai ad, ccaia,
efcee ee eeia f effecie aagee  hibii f hee ieaci. The eiig eeach idicae
ha a ecaia aach  aagee f hee aciiie i aaed  ha he bde f f ie ih he h
a  ieac ih he aia iead f he h a  ec he. I igh f he ifeai f ga ha ae
faciiaig cac ih ceacea, i i eceded ha hi beai be ed b egiai ad fdig f
eeach ad aagee accdig.
INTRODUCTION
O
The ai f ha ieaci ih fee-agig ceacea ha iceaed baia e he a eea
decade ieaia (Babe et al., 1995; Dff ad Deade, 1990; H, 1994; Lce, 1990; Sae et al., 2000;
Sid, 1991). The e ad ae f ciabe ieaci a ide, b ae geea chaaceied b ceacea
iiiaig  aig ce ad aied ii  ha (ie, ade, bae  ee  dc) ad
aig aied  eeaed ieaci (acie, acic ad ia) ih ha. Lce (1990, g. 338), decibed
ciabii i ia dhi a  fiedie ad eeig ca. Thee ieaci a  a  ie he
iiig f fd  he aia. Hee, i hi ae, ieaci ha ie feedig ae efeed  a ciabe

SC/52/WW2
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 if feedig i b e ce f a age cia ieaci ha a ica icde ceacea affiiaie behai
dieced ad ha (ee Sce f Pae, be). Sciabe ieaci diffe iai f  ba-baed dhi
ad hae achig aciiie i e f a ea hee ieeaed fac: 1) ica fc f behai (ceacea behai
dieced ad bae() ahe ha he ba ief), 2) ica ce ii f aia  ha ad 3) ica
dai f ieaci (e aied). Thee fac ae fee deeied b he degee f habiai  aaci
 ha ha he aia ehibi (Kigh ad Tee, 1995).
The iceae i ciabe cac ih ceacea aea  be age eaed  he e eai f faied
ad cecia ga ha ide iie f ca ad i  ieac ih he aia (Sid, 1991).
Thee ga fe ead  ae ad iica ccig ha-ceacea ieaci, iia  he
e ha hae bee eed f ieia (Da, 1988). The deee f hee ga ha eed i a f f
ieie ad cecia eiai f dcee i ai egi ad he d ch a Ne Zeaad
(Caie, 1999a), Jaa (Mi, 1999) ad Aaia (Gae, 1999). Ahgh  ieacie ga ie a
dcee, cecia ga ha ffe i-ae ieaci ih icee ch a ie hae (Balaenoptera
acutorostrata) i Aaia ad ga (Eschrichtius robustus) ad hbac (Megaptera novaeangliae) hae i hei
beedig ad caig aea i Meic ad he Diica Rebic (eecie), hae a bee ceaed (Ad ad
Bie, 1998; Fi, 1997).
T dae, cieific dceai f -gig ciabe ieaci ih dcee hae bee f he fai
Dehiidae, iai eeeed b bee dhi (Tursiops truncatus). Hee, a ecei ha bee
dceed f eea ccece f ia, ciabe hie hae, Delphinapterus leucas, (Fhff et al., 2000).
Sciabe ieaci ih e hae, Physeter macrocephalus, hae a bee dceed (e.g., H, 1994; Rie ad
Bedea, 1999); hee, hee aia ae aae  habiaed  ha. A ige b (Inia geoffrensis) f he
fai Paaiidae i Sh Aeica h a ied i a -gig ha-dhi fihig ceaie a eed
(Lab, 1954). Hee, he ieaci a aae eiced  he ha ad aia baiig fd ad a 
decibed a ieacie  ciabe bed hi acii.
The a  decade ha a ee he ieaia bgeig f a eaed e f id iig caie
dcee (Fhff, 1993; Fhff, 1994; Maa et al., 1999; Sae ad Sadi, 1994, 1995). Thee ga ide
he bic ih a i  i ih, ade ih, feed ad/ ch caie dcee (icdig hie hae,
bee dhi, i hae (Globicephala macrorhynchus) ad ie hae (Orcinus orca) (e.g., Dea, 1996).
Oe, bihed d ha caed ciabe ieaci i caii ih he i he id (Fhff ad Pacad, 1995) b
hi a age deciie ad he cci  eiia.
Ieacie aciiie ih fee-agig ceacea hae ei bee egaded a a -cie f f
eceai (Dff ad Deade, 1990; Richad et al., 1995). Hee, ih he iceaig ai f hee
ieaci, e e f haae  he aia, icdig eee ijie ad ee aiie, hae bee
beed dide (Ba, 1994; Ddii et al., 1995; Lce, 1990; Sa, 1997). Cee, ciei, ic
ae, aage ad he geea bic hae eeed cce ab he deeei, e  ded iac ha
hee ieaci aea  be haig  a ea e, if  a, ageed dcee idiida ad ai (Ba,
1994; Ddii et al., 1999; IFAW, 1995).
Cie a ide ih eec  he degee f egiai eaced  ec ceacea f ha acii (e.g.,
Ca, 1996; Caie, 1999a). The U.S. ha ided ha a be he  ceheie defiii f ceacea
haae  dibace. I he 1994 aede  he U.S. Maie Maa Peci Ac, haae a defied
a A ac f i, e,  aace hich (i) ha he eia  ije a aie aa  aie aa
c i he id  (ii) dib a aie aa  aie aa c i he id b caig dii f behaia
ae icdig, but not limited to, igai, beahig, ig, beedig, feedig,  heeig [ehai added].
Thi defiii a f a bad ieeai f haae  icde a behaia chage aibabe  ha
acii.
Reeach, aagee ad egiai eaed  dig ad aagig hee ieaci hae agged fa behid
hei eai. I e cae, aagee ad egiai hae  bee ieeed fig ha ijie
ificed b dhi ih h ee ee ieacig (e.g., Fhff et al. 1996; Sa, 1997; S. Jh 1991). Sch
ijie hae icded be be, bied ad ed iea ga ad bie, a f hich hae bee ei
egh  eie hiaiai. I e cae, a ia bee dhi i Bai ied a ie h a eed
abig he aia (Sa, 1997). Siia, aagee eff hae a fced  ecig he aia, eecia
i cae hee hica ha ha bee ieia ificed (e.g., Ba, 1994). Hee, age de  ac f fdig,
aagee gideie ae ica diffic  iee  efce i a egi, ee he a ei ha
ecifica adde  ee hibi he ieaci (Heig et al., 1995). Fdig ha a iied eff 
cieifica d hee ieaci, aica hgh gidia die (IFAW, 1995).

SC/52/WW2
4
La   a ac
Sciabe ceacea ica ehibi   aig degee f habiai  ha acii, b a deae a
cea aaci  ee ad bjec (ch a ba) aciaed ih he. Habiai ad aaci ae bh cideed
eaed ee (Kigh ad Tee, 1995). Habiai ca be defied a a aig f ee  a eeaed  gig
i ha i aciaed ih eihe a iie  egaie ead, eig i he i haig a ac f igificace
 he aia (Eibe-Eibefed, 1970; The, 1963). Aaci ha bee defied a he egheig f a iie
aciai ih a eeaed i (Kigh ad Tee, 1995). Aaci aife behaia a a iceae i a
aia iie aei (ia, acic,  acie)  a i. I ha bee deaed eeaed (e.g., ih fd
iiig f idife) ha aaci f idife  ha acii ca be haf  bh ha ad idife (Kigh
ad Tee, 1995).
I addii, deeiiai ad eace (bh ibe fci f habiai (Kigh ad Tee, 1995;
Richad, 1995)  ha acii ca a be haf  idife. Deeiiai ca be decibed a a deceaig
eiee  a i e ie. Teace i aifeed b eaiig ea a i egade f i egaie
igificace  he aia (Kigh ad Gie, 1995; Richad et al., 1995). Thee ee a be haf 
ciabe ceacea becae he a e i aba ad iaiae ea ee eaie  he dage ha
he ii (ch a ba ) ee. The a a be haf becae hee a be egaie higica effec
(i.e. e) accaig hee behaia ee (ahgh he ee f hee effec ae  ), eecia he
ee  he ii i chic (Kigh ad Ce, 1995; Richad et al., 1995).
R  c
N-cie idife eceai iai aie f cie aciiie i ha he iei f he e i
he fe cae i   ieia ha  ee he age aia() (Be ad Sa, 1985; Dff ad Deade,
1990). Hee, i i cea ha aciiie geea egaded a -cie, ch a idife ieig, ca ee
egaie g-e iac  id aia, egade f beig ie. Thi i aica e he ha hae a high
ga ieai ad eecai egadig hei idife eeiece (Dff ad Deade, 1990; Kigh ad Gie,
1995).
F he e f hi ae, i ae ideified a ce f ciabe ieaci ha iceae he aia
ceibii  g-e egaie iac (ee be). The ce f iac  dcee ca be he e f a)
ha aci (diec aci, ch a chaig dhi  idiec aci, ch a degadig habia ai  aiig
dieae), b) aia aci (ch a ce aachig ba   ie)  c) a ieaci beee he 
(ch a aia igeig fd  bjec ffeed b ee).
Short- and long-term responses and impacts
Ree f, ad iac , idife ae fe decibed iechageab ad ae geea chaaceied a beig f
h-  g-e dai (Fida, 1997; Kigh ad Gie, 1995). Behaia ee diec ad iediae
aciaed ih a i ae geea egaded a h-e ee. I geea, h-e behaia ee ae
cideed fa e eadi deecabe ad cecabe  iac f ha acii ha ae g-e ee (Dff
ad Deade, 1990; IFAW, 1995).
Hee, g-e ee, he he ca be ideified ad iaed f ai he aiabe, ae geea
e diec eaed  g-e iac (IFAW, 1995). Lg-e ee ae chaaceied b iac : a) hica
cdii (bi ba ie, ecgica iac,  ahgica idica), b) caie  gig behaia ae
(e.g., habiai, eace, eiiai), c) dcii ad fie (idiida, g ad ai ee) ad d)
habia e ad diibi (bed h-e aee f aia eaie  dibace) (IFAW, 1995; Fida, 1997;
Kigh ad Gie, 1995). If e f hee ae cied, edced,  aeed, e ca ae a egaie iac 
ceacea.
The eaihi beee h-e ad g-e ee i ae cea  ccie (Dff ad Deade, 1990;
IFAW, 1995). Sh-e ee d  diec aae  h-e iac; he effec f eeaed iac 
h-e behai ca e i g-e chage  behai ad hig ha effec g-e ia ad fie
(Kigh ad Gie, 1995). Fhee, hee aea  be a aea f ea beee h- ad g-e
behaia ee. F eae, h-e behaia ee ca dee i g-e behaia ae 
cdii, a he eeaed, h-e ee  a i dee i a g-e behaia chage  a acii,
ch a eiiai (he gada iceae f egaie ee  a i) (Kigh ad Tee, 1995). Sh-e
ee ca a dee i g-e, chic, behaia ae ch a eeie (Defa ad P, 1980;

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Geed, 1977), ef-decie aci (Ni ad Pec, 1961) ad eceie aggeiee (Cae, 1982) i
dcee.
Stress and control
Se a bee defied a a eiea effect [ iac]  a idiida hich eae i
[higica] c e ad edce i fie  aea ie  d  [ehai added] (B ad Jh,
1993, g. 72). Se ha a bee defied a a condition i hich aba  eee adje  behai 
hig ae ecea  ce ih he eie (Fied, 1980). The e response i a adaie ee ha
ehace he ieihd f ia i he h-e. Hee, he ieie effec f he he
ciig he e ee a hae deeei [g e] effec if ee  he e i ged
(Wab, 1991, g.339). Seie, higica chage hich ae efeed  a e a acca bjecie
ae f ffeig, b  aa (Dai, 1980).
Ha ieaci ca iac ceacea he ha ee c e aec f he aia eie ha
heie d be i he c f he aia. We-beig ha bee defied a a fci f a aia' abii  c
i eie ad  edic aeie ee (Wieea, 1985). N  ca a ac f c ead  e i a
gai (Sa, 1990), b i i e f he  ef ee ha a gai ca ece (Tae, 1987).
Iaiae behaia ee ha a be decie  he aia  i aciae a be he e f a eed f
e c i he eie (Hghe ad Dca, 1988). Pidig dhi c e hei eie ha bee
eceded a a ia ehd f iig hei efae i caii (Mai, 1991) ad a ie a 
fee-agig ceacea h ieac ih ha (Fhff ad Pacad, 1996; Fhff et al., 2000).
Baa a  ac
Becae f he iceae i ahgeic acii iig ceacea, hee i a iceaig eed f bjecie
eae f egaie iac (dibace  he f f e). Behai ha bee ed  ae he iac 
ceacea i a aie f ce, icdig haeachig, aicaf ie, ffhe i ad ga eai ad icebeaig
(IFAW, 1995; Richad et al., 1995). Sh-e behaia eaci ha hae bee decibed a ee 
ahgeic aciiie icde, (b ae  iied ), chage i: facig, eiai ad die ae ad ae;
acig ad chei f d; iig eed; ce ad ieai eaie  i; cicaie dia (e.g.,
aeia behai, caiai); acii ae/bdge ad behaia ae.
Behai i fe e f he  aaiabe idica f he cdii f aie aa ad i a ecea
ce i he eaai f hei efae (Dieaf, 1990, Seee, 1990). Meaee f higica e a be
diffic  deiabe  bai becae f cce ab addig  a ef cdii, eecia i ceacea
(Seee, 1990). Ee hica ad higica daa a  be ef he beed i cjci ih behaia
aaee (M ad Giffih, 1985). Pchgica ii ca dce a ch,  ee e, f a e ee i
aia a ca hica e (Wab, 1991). Recgii f e-eaed behai i aia i ciica 
ideificai f eahgica  ahgica cdii, hich ca cc ee if he aia aea  be i heie
ia hica cdii (Mbeg, 1985; Seee, 1990).
Hee, ee he bigica ae ae baied, a adadied, aiaie ehdg f aaig effec
f e  ceacea i acig (Dieaf, 1990). Lie bigica ee  ii, behaia ee a
ieecifica, ad ee beee idiida (Dai, 1980, 1990). I addii, i ca be diffic  deeie hehe
a behai i a diec ee  a i  i  ee  a a aciaed ih he i.
Cee, behaia aee f e i aia eie beai f behai eaie  he
eiea ad behaia ce() i hich i i beed (Hedige, 1964; Ra, 1988; Fae, 1989). Sice
dhi behai aea  a a a fci f ce (Jh ad Ni, 1986; Saaa ad Ta 1979; Wig et
al. 1990), e behai hae bee aciaed ih iea ae ch a e  dibace. Kedge f a a
eeie f behai (bh i caii ad he aa eie) aciaed ih ef cdii  iea ae
ide a aabe   ideif behai ha ae idicaie f e.
Sdig behai eaie  ceacea ecie ad ce ha eed i he ideificai f e behaia
aciai. Dhi ea ad aggeie behai hae bee fee beed gehe dig ieaci ih
ha a e a ih he dhi (Cade et al., 1970; Fhff, 1994; Ni, 1967; a, 1991; Saaa ad
Ta, 1979). Eceie ea behai ha a bee aciaed ih bed i caii (Seee, 1990). Agiic
behai, agig f aggei (dieced ad he dhi ad ad ee)  bii, ha bee aciaed
ih ai ce f dibace ad e (Saaa ad Ta, 1979). Sbiie behai ha bee aciaed ih
iiidai (Seee, 1990) ad higica e (Mbeg, 1985). Abe ha a bee ideified a a
chaaceiic f behai idicaig aggei, bii, aa  dibace (Bae, 1974; Ei, 1985; Fhff

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1993; Hedige 1964; Sae ad Sadi,1995). A ih -ca behai, ca behai a a be ed a a
eae f dhi efae. Vaiai i caiai aea  be eaed  a dhi' higica  chgica
ae (Cade et al. 1970; Fhff et al., 1996; Sjae ad Sih, 1986). Cee, ab, ca, ea ad agiic
behai hae bee ef aaee i aeig e i dce.
I i ia  e ha he abece f a eceied ee ceai de  ea ha a egaie iac did 
cc (M ad Giffih, 1985). Of ce, a ac f ee ca idicae ha he aia i affeced b, habiaed
, deeiied   ea f he acii (Fida, 1997). Hee, dibace eaci  iac ca be eai
eed f a aie f ea ha icde: a) icec ieeed idica f e, b) he eeache eece
affecig he aia befe he eeache ca ee deec he eece f he aia  bee i behai ad c) a
iabii  deec eaci becae f he be  gada deee f he behai  de  iaied iibii
(e.g., diace  ae cai).
Obc
The bjecie f hi ae ae  ide: 1) a eiia eie f ieae ha dce dcee behai
eia idicaie f e ad dibace aciaed ih ciabe ieaci beee fee-agig dce ad
ha e he a ce, 2) a eiia eie f ieae ha dce i  he aia ied i hee
ieaci ad 3) a eiia aee ad dici f hee fac i igh f aagee f, ad eeach ,
ciabe dcee-ha ieaci ad  ae ecedai accdig. Thi ae de  ide a
ehaie ieae eie  a hgh dici f he ieae. Iead, i ide a dici ad aai f ha
a be cideed a eeeai f bihed dceai  e-eaed behaia eaci ad h- ad g-
e i  dcee dig ciabe ieaci ih ha.
A eie f he dceed i ad behaia ee f ciabe dcee aciaed ih ha ieaci
i a ia e i ideifig, ieeig ad aeig he ea iac f hee aciiie  he aia. F
eeache, edge f  dibace eaci ad i i ia f hhei eig, ideificai f e
ad i aaee ad eaai f eiea ad cia aiabe i fe eeach. F aage ad ea,
ideifig ad deadig hee aaee, a e a he ai ce i hich he ae   cc, i ia
f ie-ecific aagee f iai i hich ai cdii (e.g., ecie, habia ad e g) ei. Thi
eie ad dici a ide a caie eecie f ha ha bee dceed ab he h- ad g-e
effec f ha ieaci  ciabe dcee ad eced dieci f fe eeach ad aagee i
hi aea accdig.
Sc  a
Siai i hich aia hae bee fced  be i ce ii  ha (e.g., adig, a e-eie e
fihig eai  idia aciiie) ae  addeed i he dici f hi ae ad  i he ieae eie.
Thi i becae he ciei f ciabe eie ha he aia iiiae  a he ii  ha, a ed 
i beig fced  he.
A aed eaie, ciabe ieaci a ie he iiig f fd  he aia. Hee, aciiie ch
a ceaie fihig ih ha (e.g., P et al., 1990) ad ce aciai ih fihig ba (e.g., Fe ad Wig,
1993) i  be icded i hi eie. Thi i becae hi defiii f ciabe ieaci de  icde
ece i hich i i eide ha he aia feedig i he e  ecie chaaceiic f he ieaci. Hee,
i a ca be deeied hehe feedig i b e ce f a age cia ieaci ha a ega icde
ceacea affiiaie behai dieced ad ha; ee if hee behai ae idiec eaed  baiig fd.
Cee, he ieaci i hich feedig i b e ce f a age cia ieaci (ch a iai i
hich dhi ega egage i affiiaie hica cac ih ha) ae icded. Addiia, e f ee
beig aied  eced (e.g., hed  he) b ceacea (e.g., Bade, 2000; B, 1991; Da, 1988; Ree,
1994), ee  icded i hi eie if he e cied iai f hi aci.
A eied eaie, die f he effec f ba acii  dcee (ch a ha hich cc dig dhi-
achig cie) ee  cideed eeeaie f ciabe ieaci if he acii a cied f ieaci
beee aia ad he ee  he ba a ed  ieaci ih he ba heee. O he i ied
ee ieacig ih aia f ba,  ba eig a af f hich ie eeed he ae, a
baig acii cideed a ce f ciabe ieaci.
Thi ae de  ae  ide a hgh eie f hi bjec becae  a iace f ciabe
ha-ceacea ieaci hae  bee dceed i i ad, he he hae, a edeace f he acc ae
aecda. Fhe, hi ae de  ae  eie  eaae a eia beefi ha igh be iced b
ceacea a a e f hei ieaci ih ha. La, i becae a eee f i (ch a hica cac

SC/52/WW2
7
 feedig) a  ied f a aica efeece, i de  ea ha hi eee f i a  a ce f he
ieaci;  ha i a eihe  eed f he ieaci i ha,  a he eieed bicai.
REVIEW
Pbicai ha dceed fee-agig dcee behai eia idicaie f e ad dibace
aciaed ih ciabe ieaci ih ha e he a ce ee ideified iai f a geea ieae
eie f ciabe ieaci beee dhi ad ha (Ddii et al., 1999). Pbicai ha ided he 
ee-eieed, eaic ad deaied dceai ee eeced e a bicai heee ibe.
Hee, a bicai ee icded i he abece f cieific dceai  he he fe ided
eie ifai. Cee, hi aci de  ide a ehaie eie f hi ic.
Review of stress-related behaviors
T eie behai ha ee eia eaed  e i he ieae, hei eed ccece eaie  each
ce ih hich he ee aciaed a ed f each bicai. Eff ee ade  aid iig he ae
ccece f each behai e ha ce he eied eeaed i he ae dce  he efeed  i e
ha e ce. Seeci f hee behai a baed, i a,  a ei eie f dcee behai ha had
bee eaed  ef ii  ieeed a idicaie f e i bh caii ad i he id (Fhff, 1993).
Behai ee eeced if he ee eihe ideified b he ah f he ae a eia eaed  e 
dibace  decibed iia eehee i he ieae a aciaed ih dibace (e.g., P ad Shaebege,
1991; Seee, 1990).
Ieeai f behai i high ecie- ad ce-deede. F eae, behai ch a fe-a
agai he ae, jig e ie, hig ba,  hig f bae had a be idicaie f a aie f
iea ae, icdig a, eciee ad aggei (e.g., Heig, 1993; Webb, 1978a). Cee, hee
behai ae ied ee a eia idica f e  dibace.
Obeed behaia idica f dhi e  dibace (e.g., aidace) ee ed ahe ha he eia
ce f he e  dibace (e.g., haae), egade f he eia i  beai f hica ha
ied (hi i addeed ae a Ri). Hee, hee fac ae ied (de Ce) if he ee eied a
a f he ce i hich he e-eaed behai cced i he ieae.
F each ideificai f hee behai, he fig a ed: 1) Se-eaed behai - Behai ee
decibed iai i e f hica f heee he ieae ided egh ifai  d . Hee, i
cae hee  a ieeai  eed fci f he behai a ided, hi a icded a ch; 2)
Ieaci chaaceiic - The cia gig f he aia ehibiig he behai a ed a G  Sia i
efeece  hehe he aia a ica beed ih e  e cecific  hehe i a iai a ia
aia. Thi caeg a a ecie. The e f ha ideified a he i ad/ eciie i he
ieaci f ha aica behai ee caegied a: a) Sie (icdig die ad ade), b)Bae
(icdig ee  ie be, ai aee ad he faig ), c)Feede (ie, ade, bae, 
ee f a dc idig fd  bjec  he dhi f ibe igei) ad d)Dc Pee (ee  dc
ad ie). 3) Secie f ceacea, 4) Gede f ceacea, 5) Lcai f ceacea ad ae b hich he aia a
efeed  b ee, if aicabe, 6) Ce - Eiea  behaia ce f behai, if dceed ad 7)
Ieeai  ce - ided b he ah.
The Ieaci Chaaceiic, (2 abe), f ee ee  a ecie ad aed f eea diffee
caegie f ee eaed  e behai. A, i he ieae, eie he ha caegie ee  cea
ideified i efeece  a aica behai. Cee, eea diffee ieaci e cd be ied f he ae
behai if e ha e e f ha aeaed  be aciaed (e.g., G/Bae ad G/Sie) ih a
behai. Se-eaed behai ee caified accdig  iiai (Tabe 1). The feec i hich each behai
e a eed i he ieae a aied (hee, feecie d not efec he aca be f ccece f he
behai heee).
Review of risks to odontocetes
A ecda bjecie f hi ae i  eie eee f i ccig dig,  diec eaed , ciabe
dcee-ha ieaci a dceed i he eieed ieae. F he e f hi ae, i a defied
a he aia iceaed ee  he chace f beig ijed  ied i he ce f ciabe ieaci. Thi a
iai chaaceied b hica cac ih ha  ba  ii  ha ih he eia f hica
cac (egade f hehe hica ha aca cced). The i f ii a eeded if bjec ee h 

SC/52/WW2
8
h a he aia() (egade f hehe he aia ee aca hi), ee f a diace. I Tabe 2, i ee
ideified egade f hehe   he ee ideified a i b he ah f he dce ad egade f he
eed ee f he aia()  he ee.
F each i dceed, he fig a ecded i Tabe 2: 1) Ieaci e  Sae a abe (i Tabe 1)
ad 2) Ri  aia  Caegied a f:
a) Nea Ejec  Aia i he age f ha aiig a  aca hiig  hig ih -fd bjec (e.g. a,
ea  g).
b) Nea  Aia i ce egh  ha  a f hica cac beee he , b e i ade.
c) Phica Cac (icde Nea) - Peia haf iei (e.g., a, hi, jab, e)  cai f ch
(eiie  eabe bd a). (If hica cac a ee eed f a ieaci i he ecified
dce, ee if  a a ae  eceia ee, he i a ied de Phica Cac iead f Nea.)
d) Fd  Objec Piiig  Peeig fd  bjec  he aia (icde Nea) Peia f igei f
haad ie (e.g., iaiae  ieia  ieia caiaed fd  iaiae bjec)
eeed  he aia (de  i Phica Cac e hi efeece i a ied i hi caeg).
RESULTS OF REVIEW
A a f 59 dce (icdig 19 iig ia aia) ee che f aai baed  hei caiig
ifai eaie  e-eaed behai  i. F hee, a a f 17 dcee ecie ee eed, 15
ihi he fai Dehiidae, e ihi he fai Mdidae (D. leucas) ad e ihi he fai Physeteridae (P.
macrocephalus).
S-a ba
Tabe 2 i 38 e f e-eaed behai ha ee beed i he eie. A a f 177 e f e-
eaed dcee behai (a ed  aca beed feecie) ee beed i he ce f ciabe ha
ieaci i he eie. Of hee, 107 e ee ade f ia aia. Eigh ee (a 50%) f he a
e ied ie  ad 67 (a 40%) f hee ied ia aia. Ahgh hee be d 
efec beed feecie f he aca ccece f he behai, he high eceage f e-eaed behaia
ee eed f ie ae idicae ha he iac f ie  dcee, ee i he abece f he
e f acii, a be ei de a ide aie f cdii.
Of hee behai, Aid a, b fa, he  fee eed behai, eied 31 ie. The 
fee eed he behai icded, Sa ad Sahe agai ae (eied 17 ie), fed b
fcef Phig ie (eied 14 ie), ce fed b Bie ie (eied 13 ie). The e
e f iceaed aggei beee cecific eaed  ha ieaci a eed i he ce f dhi
feedig (Oa et al., 1996). The fig behai ee eed  i he ce f ia aia: Ae 
bie, B, b,  a, Chage, Cice, Cfed, Defecae, Hi fcef, J e, Mh bd a ad
Rae. Wih he ecei f e cae f aae haae f a i hae h ae ijed a ie (Shae ad
Tee, 1993; Shae, 1995), a f he aggeie behai iig hica cac (ad a f he ijie iced b
ha) cced ih ha aeaed  be habiaed ia aia  g f aia ega iied b
ee.
M eeach a age bee eiced  die f h-e iac  he aia ece, i ae iai i
hich ideificai f diec, g-e behaia iac cd be ideified. The fig behai ee cideed
idicaie f g-e iac: 1) Aeai f fagig ae ad abiiie a e a chage i habia ad diibi
ehibied b fd-iied dhi (IFAW, 1995; Wi, 1994, 1996) ad
2) Caie ad g-e chage i behaia ee  ha acii (e.g., ibe eiiai a ed b
Caie, 1999b) ad e iace f habiai ad aaci (e.g., Heig 1993; Lce, 1990; Sae ad
Bejde, 1998) f a caificai f aia (e.g., ia, g, iied).
R  aa
A cbied a f 114 dceai f i ee beed ad ae ied i Tabe 2. Si-fie (65) ee
beed f ia aia ad 49 ied aia g. A eied eaie,  a caegie ee a
ecie  e efeece cd be eed i   e caegie. A, eie ie efeece  he ae
aia() ee eed). Thee dceai ee caified a f:
a) Nea Ejec - Eigh dceai; i ied ia aia,  ied aia fd i g (bh f
h ee ega iied ih fd).

SC/52/WW2
9
b) Phica Cac  Fif-e (51) dceai; 40 ied ia aia ad 11 ied aia g.
c) Nea- Thi- (32) dceai; fie ied ia aia ad 27 ied aia g.
d) Fd  Objec Piiig ( eceai icdig igei)  Te-hee (23) dceai; 14
ied ia aia ad ie ied aia i g.
M he e f ia aia icded ae  ii he ih fih hich a fe igeed
eidica  ih egai. Seie, feedig ee fed he bai f he ieaci i  bee ciabe
ieaci (e.g., Wai i Da, 1988). Hee, a f he ia aia ica did  aca ige
iied fih ad he he did, i a fe  fig a eid f ciabii ih ha (e.g., B, 1991;
Lce, 1978; Lce ad Mi, 1986), he he aia ee ijed (Ddii et al., 1995), ega (Wai i
Da, 1988)  he e g (Cii et al., 1998).
N-faa, ieia ac f aggei dieced ad aia icded f e f ia dhi a
f: Ndg a hi ad caed f a ea (Bge, 1983), e bjec icdig fihig gea ad c ee
h a Wia ad K (Fhff et al., 2000) ad Dad a h a ih a g (Lce, 1978). Ne
acc f -faa, ieia ac ee a dceed f aia ihi g, a f hich cced ih ega
iied dhi: Ba (1994) eed he da fi f eea dhi beig c ff; acc f dhi baied
ad hed, e icdig ae  ad he; ad dhi fed ba, i ad fie cace. Sae ad Bejde
(1998) eed a dhi beig hi i he face. A, e -faa, -ieia iac f fihig gea (e.g.,
B, 1991; Ba, 1994), ach (Sae ad Bejde, 1998) ad ee ad ba (Ae, 1963; Ddii et al.,
1996; Fhff et al., 2000; Lce, 1978) ee eed f dcee i he ce f ciabe ieaci ih
ha.
Ne aiie hae a bee aciaed ih ciabe ieaci ih ha. Faaiie f hich he
eed cae f deah a ieia ha icde a ea f dhi; hee ia dhi, BW, O, ad
Ca Rica ad e ega iied dhi beed ihi a g, Od Chaie (Da, 1988). Ne -
ieia aiie, a aciaed ih dhi feedig, hae a bee eed i he ce f ciabe ha
ieaci. Thee icde a e f ee dhi h ee beieed  hae died f i-eaed i ad
iceaed caf ai ied  fd iiig (Ba, 1994; IFAW, 1995; Wi, 1994, 1996). Ne ad
ici diaeaace f he aia hae a bee ed, fe diec afe he aia had bee baed a he
ce f aace  cfic (Da, 1988).
Rac a aa
Seea eeache ed diffic i iaig he effec f fd iiig f ba acii ad ba acii
f ie acii (e.g., Caie, 1995; Fhff ad Ddii, 1996; Sae ad Bejde, 1998). A, ih fe
ecei (e.g., ie dhi i Haaii; Wig, 1999 ad d dhi i Ne Zeaad; Ba, 1997; Yi, 1999;
Wig et al., 1998), e-iac baeie daa f  dcee ai a acig f  ageed ai.
Mch f he eeach cied f deciie acc f iica ceced beai (e.g., Da, 1988).
Hee, he fig bicai dce eaic beai ad aiaie aai f e-eaed behai
i fee-agig dce ied i ciabe ieaci ih ha: Ba, 1997; Bejde, 1997; Bejde et al., 1999;
Cb, 1999; Caie, 1995, 1999a, 1999b; Caie ad Bae, 1997; Fhff ad Ddii, 1996; Fhff et
al., 1996; Fhff et al. 2000; Gffa et al., 1999; Heiich-Ba et al., 1994; Ra, 1998; Sae ad Bejde,
1998; Wd 1999, Yi 1999.
The ieae eeaed ha he fc f eeach  aia g a  fee he behai f he g 
d (e.g., ceai f ieaci, d diea) ahe ha ha f a idiida fca aia (e.g., fe-a, e-h
hea). T e degee, hi a ifeced b he beai af beig ad-baed (e.g., Bejde et al., 1999), ba-
baed (e.g., Caie ad Bae, 1997), a e f deae (e.g., Fhff ad Ddii, 1996) Vai adaage
ad diadaage f diffee eeach af ee ed. F eae deae ad ba-baed eeach eia
iaced aia hie ad-baed eeach did  ei ecei f e be  deae behai. M
die  beed -ca behai eaie  ha acii (ih a abe ecei; Scaaci et al., 1999).
Maagee ga ee geea decibed a diffic  iee, eecia he egiai a  eaced 
 hibii  gideie (Heig et al., 1995). Hee,  f hee ga ee cideed a ea
deae ccef ce he ee ieeed (Fhff et al., 1996; Fhff et al., 2000; S. Jh, 1991).
DISCUSSION
S-a ba

SC/52/WW2
10
I i  iig ha Aid ad Sa ad Sahe ee  fee ideified a aaee f e 
dibace. Aidace eaci hae bee cideed e f he  bi aifeai f dibace (Richad
et al., 1995) ad ahe agai he ae face ae high iibe f ad  ba eaie   he ceacea
behai.
Interpreting behavior
Ieeai f behaia ee a cea ce- ad ecie-deede. F eae, ahgh aidace
a fee ideified a a dibace ee, i a a ccaia decibed a a aec f a behai (e.g.,
Da, 1988). Behai ha ee ieeed i e ce a aggeie (e.g., fe-a, fcef bd  ba cac,
hig bd a), ee ieeed a eciee, a,  ieigai i ahe. A, a chage i behai ee
 eceai ieeed a idicaie f egaie iac (e.g., Bejde et al. 1999; IFAW, 1995).
The ieeai f aggeie behai a a idica f e i ce ad e eeach i hi aea i
ceai aaed gie he feec i hich habiaed dhi ehibi hi behai ad ha. A ih caie
dhi (P, 1973; 1981), habiaed dcee a diec he ae diace-eaed behai ad ha ha
he d ad cecific ad a eec he ha  ecicae a a cecific d. Thee aia a
eec ha  fi he cia e a ccied b cecific (a hee decibed a aiiai
edec i he aai f ieecie ieaci i caii [Hedige, 1964]). Thi a acc f he ccece f
ea ad aggeie behai dieced ad ie i acciaed/habiaed dcee. Hee, e eeach
i eeded  deeie ha a ad eceie ee f aggei ae i he ce f bh ceacea-ceacea a
e a ceacea-ha ieaci.
Related examples
Ma f he behaia ee eed i he ieae hae bee beed i he ce i hich dcee
ee i ce ii  ha. F eae, dhi cagh i a e-eie e i ce, i-ae, ii 
ha hae ehibied chei f dhi g; e-h hea ad head; fe ad bd a ha ee ieeed
a agiai; aace; aa; aig ad hea (Ni ad Dh, 1980; P ad Kag,1980; P ad
Shaebege, 1991).
Sdie f he behai f caie dhi ed i i-ih-he-dhi ga hae a dceed iia
behai ha ee ieeed  be idicaie f agiic ee  ie. I a aee f hee ga i
he U.S. (Fhff, 1993; Fhff et al. 1996) ad i he Bahaa (Fhff, 1994), he fig eea behai ee
beed: Vaiai f hie e ad ae, ab a agai ae, ea fcef behai, ae  bie, bie ad
aidace. I a d f hee ga i he Uied Sae, Sae ad Sadi (1994; 1995) ed he fig
eea behai: Oe-h hea; hi; fich; fee; ab-; aid aach; ja-ca; bd-a; h; head-je;
ai-a; ab b ad ea behai.
I eig/feedig ga ha ie caie dcee ieacig ih ee a he edge f ece,
aidace, ccef ae  eea ad dii f ee ee beed (Maa et al., 1999). A, e
behai eed dieced ad aie i he ae hae bee beed i caii ad ce aae he
beed i ia, ciabe dcee. Thee behai icde chagig, big, biig, gabbig, dig ad
hdig he aie  he b f he a (Defa ad P, 1980; Seee, 1990). Cee, a a beed f
ia aia, i i  c f caie dce  ije he ee ih h he ieac.
S- a - ac
The ieae ide eea eae f he high e eaihi beee h- ad g-e iac i
dcee. The h-e iac a be ea  iie b he iae g-e iac a be egaie. F
eae, dhi hae ehibied iie h-e ee (b ehibiig aaci)  ee iiig he ih
fd, ee hgh i ha bee deaed ha fd iiig ie eed i iceaed dhi ai i hee
iai (IFAW, 1995; Wi, 1994, 1996). A Bejde et al. (1999, g. 748) aed, O iei i ha he aji
f iac  biic eie caed b eci ee, ch a dhi-ieig eai, ae caie
ahe ha caahic.
Fhee, e ia (Fhff et al., 2000; Lce, 1978, 1985) ad fd-iied aia (e.g., Ba,
1994), did  ehibi a egaie behaia ee  ii (e.g.,   hica cac ih ha) ha had
e iji  he ae aia i he a. I e cae, he aia ehibied a cied iie aaci 
he ii deie e ijie iediae iced a a e. Thee ee a be idicaie   f
habiai, b f deeiiai  eace (Kigh ad Gie, 1995; Richad et al. 1995) ad iae he
dage ha hee behaia cee e  he ia f idiida  ai de hee cdii.

SC/52/WW2
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CONCLUSIONS AND RECOMMENDATIONS
Oea, hi eie idicae ha ee he  e-ieied ciabe ha ieaci ih dcee ae
accaied b a  be f edicabe iac ad i  he aia, e f hich ae g-e ad
ife heaeig  idiida a e a  ai. I a deae ha dce ehibiig he highe degee
f cac ih ha (e.g., he habiaed ad aaced  ha) ae geea a he geae i f ij, ie, 
deah. I aica, icide i hich ha ieia ijed  ied dcee ied i ciabe ieaci
ee eed a ecie f ia aia ad aia ega iied ih fd. Hee, a
dibace eaci i dcee h ae  a aaced  ha a be eed i becae he aia ae
geea  a iibe. Fhee, becae f he a difficie ihee i beig behaia iac 
dcee ed i he ieae, hi eie idicae ha a deeiai f   iia iac  ceacea hd
 be ade baed  he abece f eceied h-e ee ae.
F he eie, i i a cea ha ai fac f ha acii iacig dcee deeie, i a, bh
he iei ad age (h-  g-e) f he aia behaia ee (Kigh ad Ce 1995). Thee fac
icde: a) ageeabii  he aia; b) cabii ad edicabii f he acii b he aia; c) iig; d)
feec; e) agide; f) cai ad diace f he aia ad g) dai (e.g., IFAW, 1995; Bejde, 1997;
Caie, 1995; Fhff, 1996; Gee ad Cae, 1999; Ra, 1998).
Cee, he effec ha hee i fac hae  he aia behaia ee i a deede 
ai chaaceiic f he age aia() (IFAW 1995; Kigh ad Ce, 1995; Richad, 1995). Thee fac
icde: a) ecie; b)behai a he ie f he aach; c) hi f ad ei ee  ee  he ae 
iia i; d) age, gede ad edcie a; e) habia ad eiea cdii; f) g ie ad
cii; g) higica cdii ad h) ie f da ad ea (e.g., Ba 1994; Caie, 1999a, 1999b;
Caie ad Bae, 1997; Ra, 1998; Rie ad Bedea, 1999; Wig, 1996; Yi, 1999).
The cei f he eaihi beee he chaaceiic f ha acii ad he chaaceiic f he aia
ad i eie a eide ac die f diffee ieaci. F eae, he beed iac f e
ha ie a eed iia i e eie (e.g., Bejde, 1999; Ra, 1998) hie he iac f a
ige ie i he eie (e.g., Dic-Lid ad a-Lid, 1999) a eee egh  e i
beai f eeaed dii f e ad he ia aciiie. Fhe eeach i he eaie effec f hee
fac a e a he caa eaihi beee hee h-e behaia ee ad g-e iac i cea
aaed.
The e iace f behaia idicai f e ad dibace i he ieae idicae ha behai i
fe he  beabe ad eaabe aifeai f egaie iac  dcee. Hee, i i cea ha
behaia aaee f iac  dcee eed  be ehaie ad cie beed, ideified, aified
ad aaed i cjci ih e bigica ad eiea aiabe befe deeiai f acceabe
ee f iac ca be ade. A a ed i he ieae, each e f beai af caie ih i ai
adaage ad diadaage. Theefe, behaia aaee f idiida aia a e a ai beed
f ad, ba ad deae, he ibe, ae eied  ide a ceheie ice f ieaci ad hei
effec. A, g-e, aiaie die f he behai, big ad ecg f ceacea ageed f eceaia
ha acii hd be iiiaed a ea a ibe; efeab befe he iac,  ha h- ad g-e chage
hgh he deee f he acii ca be e eadi ied ad aeed (e.g., IFAW, 1995).
The eeach deae ha ideificai f e-eaed behai i aabe i deeiig iac  aia
e a iiiig iac he icaed i aagee ga. Hee, i eieig he ieae, i a
eide ha aagee f ieaci ih dcee, he ieeed, a fe h iadeae ad acig
aiae eeach, iig, edcai ad efcee iae  he ee ad iei f he ieaci.
Maagee aeaed  be he  effecie he i a baed  eeach, a ie- ad ecie-ecific ad a
ieeed ea i he deee f he ieaci (e.g., Caie, 1999a; Fhff et al., 2000; S. Jh, 1991). I
he eie f eeach, e beai a fee ed; ha a ecaia aach  ieacig ih ceacea i
aaed  ha he bde f f ie ih he h a  ieac ih he aia ahe ha he h a 
ec he (e.g., Caie, 1995, 1999a; Fhff, 1993; Heig et al., 1995; IFAW, 1995). I igh f he
ifeai f ga ha ae faciiaig cac ih ceacea, i i eceded ha hi beai be ed
b egiai ad fdig f eeach ad aagee accdig.

SC/52/WW2
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Tabe 1. Te f eia e-eaed dcee behai eed f ciabe ha ieaci. A ieacie
behai ee dieced ad ha e heie ed. Nbe efe  he be f ie ha each behai
e a eed i he ieae ad d not efec he aca be f ccece f he behai heee.
Nbe i aehee efe  beai f ia aia a a be f he a f each behai e.
Ab behai  2
Agiai, Aggeie, Fcef,  Rgh behai  5 (4 ia)
Aggeie behai ih cecific  1
Aid b fee, die, chage dieci,  i aa  31 (13 ia)
Ae  bie  2 (a ia)
Bie  13 (7 ia)
Bbbe eeed  1
B, B,  Ra  8 (6 ia)
Ceai f ieaci  2
Chage f behai  9 (4 ia)
Chage f ee e ie - 1
Chage  1 (a ia)
Cice  1 (a ia)
Cfi  1 (a ia)
Defecae  1 (a ia)
Diea f d -3
Fich  3 (2 ia)
Fagig iceece  1
Head d  je  3 (2 ia)
Hi fcef  9 (a ia)
Hi  a ba  bjec  3 (2 ia)
Iei f behai (e.g., e  fage)  4 (2 ia)
J e  4 (a ia)
Mi i igh g  1
Oe,  Oe-Sh h dia  5 (4 ia)
Mh bd a  4 (a ia)
Ph (fcef) ba  5 (a ia)
Ph (fcef) e  14 (12 ia)
Rae  1 (a ia)
Ri behai  1
Sea behai  9 (8 ia)
Sa  ah bd a  ae  17 (7 ia)
Sbiie behai - 1
S-h 1
Thea dia  2 (a ia)
Udeae (dag  hd) - 4 (3 ia)
Vaiai i caiai  3 (1 ia)
Ta  177 (107 a)

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Tabe 2. Ri  dcee eaed  ciabe ieaci ih ha.
R  Aa Iac T Rc, Sc1, a Na  Aa ( Sa)
Nea-Ejec Sia Dhi-
Sie,
Wade, Bae
 a Dc
B, 1991 (T, Fedd; aig h a aia); Bge, 1982 (T,Ndg; ea h ad hi aia, aia
hi ih a); Da, 1988 (D, BW; g h - ied); Da, 1988 (T, Ca Rica; h ad ied); Fhff et
al., 2000 (D,Wia ad K; K hi ih a; bjec ch a fihig ie/e, c, ba, ife jace,
ba fede, ad ae  h a aia); Lce, 1978 (T, Dad  Bea; h ih g)
Nea-Ejec Dhi G-
Sie,
Wade, Bae
 a Dc
Da, 1988 (T,Od Chaie; eab h ad ied); Sae ad Bejde, 1998 (T, Ach ded ea 
 dhi)
Nea Dhi G-
Sie,
Wade, Bae
 a Dc
Ba, 1997 (L); Bejde, 1997 (Ch); Bejde et al., 1999 (Ch); Caie, 1995,1999a,b (Dd,G,O,Pc,T);
Caie ad Bae, 1997 (T, Dd); Da, 1988 (T, Whiiaga dhi); Dic-Lid ad a-Lid, 1999
(S); Ddii, 1996 (Sf); Fe, 1999 (S); Gee ad Cae, 1999 (S); Fhff, 1996 (Sf); Fhff ad
Ddii, 1996 (Sf); Fhff ad Pacad, 1996 (T); Heiich-Ba et al., 1994 (G); Mead,1962 (O); Mi,
1999 (T, S); Pead, 1986 (S); Paa ad Mae, 1999 (S); Ra, 1998 (Sf,T); Rie, 1996; Rie ad
Bedea, 1999 (T, Sb, Sf, G, P, Md); Sa, 1999 (O); Scaaci et al., 1999 (T); Wei et al., 1996 (T), Yi,
1999 (L)
Nea Sia Dhi-
Sie,
Wade, Bae
 a Dc
Caie, 1999a (T, Mai); Da, 1988 (O, Ea; T, Wai ); Dbb, 1984 (T, Jea-Li); Peeie,
1985 (T, Jea-Li)
PhicaCac Dhi G-
Sie,
Wade, Bae
 a Dc
Ba, 1997 (L); Ba, 1994 (T); C ad Se, 1985 (T); Da, 1988 (Dd; Ne Zeaad, S; Bai);
Heig, 1993 (Sf); Oa, 1995 (T); Oa et al., 1996 (T); Sae ad Bejde, 1998 (T); Shae, 1993, 1995
(G); Wd, 1999 (T)
Phica Cac Sia Dhi-
Sie,
Wade, Bae,
 a Dc
Ae, 1963 (T, O); Bige et al., 1999 (T,Pia); B, 1991 (T, Fedd); Bge, 1982 (T, Ndg);
Cea ad Die, 1975 (T, D ad Nia); Da, 1985 (L, Ta); Da, 1988 (Dd, Ea; Sf,
Sad; T, Chaie; T, Ca Rica, T, Jea-Li, T, Wai); Da, 1988 (L, Ta; S, Sad; T,
Jea-Li; T, Si); Dbb, 1981 (T, Hace; T, Dbbie); Dbb, 1981 (T, O); Ddii et al.,
1995 (T, Pia); Fhff et al., 1996 (T, Pia); Fhff et al., 2000 (D, Wia; D, K); Gichi, 1967 (T,
Chaie);Gffa, 1996 (T, H); Gffa et al., 1999 (T, H); He, 1967 (T, Dad); Lce,
1978 (T, Dad  Bea); Lce, 1990 (T, Si); Lce ad Mi, 1986 (T, Pec); Ld, 1967
(T,Geg Gi); Mi ad Lce, 1988 (T, Si); Peie, 1990 (T, JJ); Sa, 1997 (T, Tia); S.
Jh, 1991 (T, JJ), Va de T, 1992 (T, Fie); Webb, 1978a,b (T, Dad  Bea)

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R  Aa Iac T Rc, Sc1, a Na  Aa ( Sa)
Fd  Objec
Piiig (
eceai icdig
igei)
Dhi G-
Sie,
Wade, Bae
 a Dc
Ba, 1994 (T);Cb, 1999 (T); C ad Se, 1985 (T); Gabe ad Gabe, 1997 (Sc);Oa et al.,
1995 (T); Oa et al., 1996 (T); Sae ad Bejde, 1998 (T); Wi, 1994 (T)
Fd  Objec
Piiig (
eceai icdig
igei)
Sia Dhi-
Sie,
Wade, Bae
 a Dc
Ae, 1963 (T O); Cii et al., 1998 (Sf); Da, 1988 (O, Ea; T, O; T,Wai); Ddii et al.,
1995 (T, Pia); Fhff et al., 2000 (D, K; D, Wia); Gffa, 1996 (T, H); Gffa et al., 1999
(T, H); Lce, 1978 (T; Dad  Bea); Ld, 1970 (T, Geg Gi); Sa, 1998 (Sf); Webb,
1978a (T,Dad  Bea)
1 Secie abbeiai ae a f:
Ch = Cephalorhynchus hectori, Hec dhi
Dd = Delphinus delphis, C dhi
G = Globicephala macrorhynchus, Sh-fied i hae
Ig = Inia geoffrensis, B
L = Lagenorhyncus obscurus, D dhi
Ob = Orcaella brevirostris, Iaadd dhi
O = Orcinus Orca, Kie hae
Md = Mesoplodon densirostris, Baiiee beaed hae
P = Physeter macrocephalus, Se hae
P = Pseudorca crassidens, Fae ie hae
Sb = Stenella bredanensis, Rgh-hed dhi
Sc = Sousa chinensis, Id-Pacific hbac dhi
Sf = Stenella frontalis, Aaic ed dhi
Sf = Sotalia fluviatilis, Tci dhi
S = Stenella longirostris, Sie dhi
S = Sousa teuszi, Aaic h-baced dhi
T = Tursiops truncatus, Bee dhi