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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.

  • Terramar Research and SONAR
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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,
SC/52/WW2. 20 .
Baa Ica  S  Oc
D Iac  Ha:
A Pa R a Dc
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
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.
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
 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
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).
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,
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 
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;
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
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
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).
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
 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.
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 
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).
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.
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
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).
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
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).
S-a ba
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
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.
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
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.
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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)
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-
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-
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-
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-
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-
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-
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)
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
Dhi G-
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
Sia Dhi-
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
... Aspects of habituation and conditioning can be detrimental to the animals, these include desensitization to potentially harmful situations and reduction in instinctual responses to avoid risky situations that could be harmful to the animals health (Dugatkin 2004). Such desensitization is evident in SSDs, as they are at the highest risk of injury, illness and death due to their high degree of contact with humans, human activities and often poor quality habitat (Frohoff 2000;Frohoff, Vail and Bossley 2005;Simmonds and Stansfield 2007). ...
... Reference Bumping dinghies (Müller et al. 1998) Tail slaps (in proximity of approaching boats) (Müller et al. 1998) Pushing swimmers with rostrum (Dudzinski, Frohoff and Crane 1995;Eisfeld, Simmonds and Stansfield 2010;Müller et al. 1998) Playing with foreign objects (e.g. balls, plastic rings) (Berrow 2009;Lockyer and Müller 2003;Webb 1978a;Webb 1978b) Circling in a small area (Lockyer and Müller 2003) Circling boats (Lockyer and Müller 2003;Müller et al. 1998) Escorting boats (Bloom 1991;Bloom et al. 1995) Approaching & nudging propellers (Dodds 2009;Müller et al. 1998) Playing & rubbing ropes, anchor chains (Bloom 1991;Bloom et al. 1995;Lockyer and Müller 2003;Webb 1978a;Webb 1978b;Wilke, Bossley and Doak 2005) Nudging & pushing boats, canoes and kayaks (Lockyer and Müller 2003;Morris and Lockyer 1988; St John 1991) Preventing swimmers leaving water (either by separation, pushing or holding swimmers/divers on sea floor) (Berrow 2009;Eisfeld, Simmonds and Stansfield 2010;Frohoff 2000;Lockyer and Müller 2003;Webb 1978a;Webb 1978b;Wilke, Bossley and Doak 2005) Dolphin breaches onto or over a swimmer (Berrow 2009;Eisfeld, Simmonds and Stansfield 2010;St John 1991;Webb 1978a;Webb 1978b) Dolphin leaps over bow of boat, surfboard etc. (Lockyer and Müller 2003) Dolphin tows a swimmer who is holding onto its dorsal fin (Eisfeld, Simmonds and Stansfield 2010) Bowriding and wakeriding vessels (Eisfeld, Simmonds and Stansfield 2010) Remaining stationary (and logging or snagging) near swimmers, boat, anchor, buoy etc. (Barbieri and Bearzi 2000;Eisfeld, Simmonds and Stansfield 2010;Webb 1978a;Webb 1978b) Towing boats (e.g. holding onto rope of boat and pulling through the water) (St John 1991;Webb 1978a;Webb 1978b) Sexual behaviours directed towards swimmers, boats and objects (Bloom 1991;Bloom et al. 1995;Dudzinski, Frohoff and Crane 1995;St John 1991;Webb 1978a;Webb 1978b) 'Stealing' objects to which may be played with (St John 1991) 'Gift giving'; dolphin presents objects to people e.g. ...
... Frohoff and Packard (1995) note that '…the only documented accounts of free-ranging bottlenose dolphins directing aggressive behaviour towards humans have occurred when the dolphins have been in highly stressful or unusual situations'. Frohoff (2000) further notes that the level of stress experienced by a dolphin during interactions with swimmers can be serious 'under a variety conditions'. ...
... The commercial aspects of observing cetaceans in their natural habitat are certainly preferable to the unsustainable (let alone cruel) aspects of intentionally slaughtering. Yet, it is critical to acknowledge that close contact with any wild animals, including dolphins and whales, can present serious risks for cetaceans (and humans) and implementation of precautions are warranted to protect them [66]. ...
... In the only on-site assessment of multiple species of solitary, sociable odontocetes (toothed whales, dolphins and porpoises) over numerous years, Frohoff identified three critical aspects of successful protection of solitary individuals (especially apparent when implemented in stewardship programs with Catherine Kinsman designed to protect orphaned and isolated belugas); (1) early assessment of the unique and varied risks encountered by each individual as well as any distinctive behavioral or physical qualities, (2) carefully designed, but nimble and quickly-implemented protocols tailored for each circumstance to mitigate risks to the cetaceans (and sometimes humans), and (3) early implementation of research (including aspects of communicative, cognitive, psychological and emotional complexity) feeding back directly into the second aspect, risk management and promoting wellbeing [66][67][68][69][70][71]. ...
... There is no single and comprehensive prescription for the design of protocols for ICR, yet some commonalities have been observed across species and situations that can inform this approach. In groups of cetaceans as well as solitary individuals who interact closely with humans, research has shown that successful efforts are directly related to early implementation and consistency of on-site, pro-active protection and research programs [66,68]. Studies on lone, sociable cetaceans of various species such as bottlenose dolphins and beluga whales [62,68,69] have shown that ongoing, systematic research integrated with responsible decisions about protection and conservation provide vital feedback on how our interactions with cetaceans affect their welfare. ...
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Abstract: Contemporary knowledge of impressive neu- rophysiology and behavior in cetaceans, combined with increasing opportunities for studying free-ranging ceta- ceans who initiate sociable interaction with humans, are converging to highlight serious ethical considerations and emerging opportunities for a new era of progressive and less-invasive cetacean research. Most research on ceta- cean cognition has taken place in controlled captive settings, e.g., research labs, marine parks. While these environments afford a certain amount of experimental rigor and logistical control they are fraught with limitations in external validity, impose tremendous stress on the part of the captive animals, and place burdens on populations from which they are often captured. Alterna- tively, over the past three decades, some researchers have sought to focus their attention on the presence of free- ranging cetacean individuals and groups who have initiated, or chosen to participate in, sociable interactions with humans in the wild. This new approach, defined as Interspecies Collaborative Research between cetacean and human, involves developing novel ways to address research questions under natural conditions and respect- ing the individual cetacean’s autonomy. It also offers a range of potential direct benefits to the cetaceans studied, as well as allowing for unprecedented cognitive and psychological research on sociable mysticetes. Yet stringent precautions are warranted so as to not increase their vulnerability to human activities or pathogens. When conducted in its best and most responsible form, collaborative research with free-ranging cetaceans can deliver methodological innovation and invaluable new insights while not necessitating the ethical and scientific compromises that characterize research in captivity. Further, it is representative of a new epoch in science in which research is designed so that the participating cetaceans are the direct recipients of the benefits.
... The absence of stress-related behavior or indicators of stress does not necessarily indicate a state of well-being, as indicators are not always apparent (Frohoff 2000) and stress may result in only slight physiological changes (St. Aubin & Dierauf 2001). ...
... Aubin & Dierauf 2001). In a paper on behavioral indicators of stress in odontocetes, prepared for the International Whaling Commission, Frohoff (2000) recommends a precautionary principle in managing guidelines for human/dolphin interactions, as stress is not always observable. ...
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An emphasis on physical aspects of harbor seal well-being in the wild and in rehabilitation within the literature leaves a gap in knowledge about their psychological well-being that is especially relevant to improving standards of care in rehabilitation and long-term captivity. While harbor seals are a common and widely distributed coastal species, some crucial endogenous traits that apply directly to assessing physical and psychological well-being are elusive and highly in dispute within the literature. For example, do harbor seals develop social bonds with one another and does a social structure exist within the group haul-outs? These questions are among the disputed topics in studies on harbor seals. I adopt a heuristic model in this study, drawing on knowledge of experienced seal researchers and caregivers through interviews, to develop an understanding of well-being, stress, sociability, and other topics. Parallel responses within the interviews highlight harbor seals’ sensitivity to disturbance and vigilance, their curiosity and accommodation to humans, the importance of play, their strong cognitive and spatial abilities, the mother/pup bond, foraging as an expression of their sensory, cognitive and spatial abilities and an certain sociability. I applied these attributes to the care of harbor seal in captive settings, and suggest that access to a stimulating and varied environment, social groups, objects for play, foraging opportunities and limits on noise and commotion all support harbor seal well-being. Five out of seven experts agreed that harbor seals live within social structures in the wild and form social bonds, and almost all agreed that solitary housing in captivity is inadvisable and could be harmful. Analysis and synthesis of the interviews and evidence from the literature contribute to a framework of well-being, presented in a table format, that can serve as an evaluative tool for assessing well-being and care in captive settings and well-being in the wild.
... La observación de ballenas con fines turísticos inició en 1 Norteamérica en la década de 1950; actualmente, predomina la observación de la ballena gris, la orca y la ballena jorobada, aunque otras especies de ballenas, así como los delfines y marsopas también despiertan curiosidad e interés en el turista. Esta actividad se ha difundido rápidamente alrededor de todo el mundo y en las últimas dos décadas el número de países, así como en el número de personas que la realizan, ha aumentado significativamente (Findlay, 1997;Brandao et al., 2000;Frohoff, 2000;Ávila y Alvarado, 1998;Malcom y Lochbaum, 1999). ...
... La información que se tiene acerca del desarrollo y las condiciones en la que ocurre la habituación en las poblaciones en general es muy limitada (Richardson et al., 1995) y puede modificarse por cambios en el ambiente que altera los niveles de estrés en los organismos (Alados et al., 1996). Así mismo, se sabe que la habituación puede llegar a ser riesgosa hasta cierto punto para los animales, en cetáceos sociales, se ha observado que la anormal indiferencia o la poca reacción de los animales al peligro (presencia de embarcaciones) resulta ser dañino para los mismos (Frohoff, 2000). ...
Influencia del turismo sobre la conducta del lobo marino de California Zalophus californianus en la lobera Los Islotes, B.C.S., México.
... This has reverberated in financially viable businesses based on taking tourists to see them, contributing to a rapid growth in dolphin watching tourism. The development of such activities has resulted in a form of intensive exploitation of these animals around the world (Simonds, 1991;Frohoff, 2000). In fact, there have been reports of harassment to the animals, including severe injuries and in some cases; mortalities have been accounted worldwide (Bryant, 1994;Lockyer, 1990;Santos, 1997). ...
... In fact, there have been reports of harassment to the animals, including severe injuries and in some cases; mortalities have been accounted worldwide (Bryant, 1994;Lockyer, 1990;Santos, 1997). As a consequence, scientists, managers, policy makers and the general public have expressed concern about the detrimental, yet poorly understood impacts of these activities on tourism-targeted cetaceans (Dudzinskiet al., 1994;IFAW, 1995;Frohoff, 2000). ...
... This has reverberated in financially viable businesses based on taking tourists to see them, contributing to a rapid growth in dolphin watching tourism. The development of such activities has resulted in a form of intensive exploitation of these animals around the world (Simonds, 1991;Frohoff, 2000). In fact, there have been reports of harassment to the animals, including severe injuries and in some cases; mortalities have been accounted worldwide (Bryant, 1994;Lockyer, 1990;Santos, 1997). ...
... In fact, there have been reports of harassment to the animals, including severe injuries and in some cases; mortalities have been accounted worldwide (Bryant, 1994;Lockyer, 1990;Santos, 1997). As a consequence, scientists, managers, policy makers and the general public have expressed concern about the detrimental, yet poorly understood impacts of these activities on tourism-targeted cetaceans (Dudzinskiet al., 1994;IFAW, 1995;Frohoff, 2000). ...
... Solitary, sociable odontocetes exhibit many characteristic behavioral patterns that may result in their injury or death or increase their risk of these due to proximity to human activity. In fact, odontocetes exhibiting the highest degree of contact with humans appear to be at the greatest risk of injury, illness and death from humans or human activity (Frohoff 2000). In particular, incidents in which humans intentionally or accidentally injured or killed odontocetes were reported almost exclusively for solitary animals and animals regularly provisioned with food. ...
... These situations have been the focus of much recent deliberation and debate among scientists, managers, and the public. Relatively few studies of these animals have been published (but see Frohoff 2000and Wilke et al. 2005); and even less information is available about their management. Thus, management of these situations is often undertaken with little knowledge of previous successes and failures involving the same or similar species. ...
... Solitary dolphins frequently seek out interactions with humans, and this behavior may provide important physical contact, substituting that usually provided by conspecifics (Wilke et al., 2005). When exposed to uncontrolled human contact, however, these interactions may have negative consequences for the dolphin, leading to injuries, lacerations, or death (Lockyer, 1978;Frohoff, 2000). ...
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Social organization is an important feature of the behavior of most delphinids, including the rough-toothed dolphin (Steno bredanensis), although, even in social dolphins, specific circumstances may lead to solitary behavior. In these situations, a solitary dolphin may shift to interacting with humans. Between November 2018 and April 2019, we obtained seven records of the occurrence of a solitary juvenile rough-toothed dolphin in Sepetiba Bay, southeastern Brazil, within an area inhabited by a resident population of Guiana dolphins, Sotalia guianensis. The solitary dolphin occupied a small home range located in an inner area of the bay, close to a port and industrial installations. This area is known to be inhabited by shoals of mullet (Mugilidae) and is not visited frequently by the resident Guiana dolphins. The solitary rough-toothed dolphin emitted three types of vocalization: pulsed calls (N = 9), echolocation click trains (N = 12), and whistles (N = 3), and spent most of its activity time interacting with humans (42%) and foraging (32%). The area occupied by this dolphin, together with its low vocalization rate, suggest the avoidance of agonistic interactions with local Guiana dolphins. The behavior pattern presented by the solitary rough-toothed dolphin indicates the third stage of development of friendly solitary dolphin behavior. At this stage, a management plan is urgently required to avoid the negative consequences of the human-dolphin interaction.
... Other critiques of human/dolphin spaces are founded on constructs that involve biological, environmental and psychological factors, noting the stresses that result from enclosure (e.g. Frohoff, 2000;Marino, Bradshaw, & Malamud, 2009;White, 2007). Key opponents (Rose, Parsons, & Farinato, 2009, n.p.) to cetacean captivity equate it with the abuse of human children: ...
Scholarship recognizes the co-construction of space by humans and non-human animals (hence, animals), but the complex geographies of some animals whose lives depend upon human care remain under-studied. This article explores human–dolphin relations within the context of Dolphin-Assisted Therapy (DAT), a practice in which most dolphins are in human care. We trace a genealogy of dolphin–human relations in built environments, and draw on a DAT case study in Curacao, to understand how the entangled agencies of humans, dolphins and other actants have co-constructed spaces of mutual therapy and care. Our research highlights the circumstances of ‘legacy dolphins’ in DAT, dolphins whose lives depend on human care. We suggest that, while the services of dolphins are recommodified through DAT, the legacy dolphin is de-commodified through ‘relations of obligation’ built on mutual ‘caring for’ as both companion species and work colleague.
... morbillivirus), environmental factors (e.g. abnormal temperatures), biotoxins, and/or contaminants, all of which may lead to decreased immune function(Fair and Becker 2000, Ross 2002, Evans 2003, Halvorsen and Keith 2008.For example, Van Bressem et al. (2009a) observed that in the case of cetacean morbillivirus in Mediterranean striped dolphins (Stenella coeruleoalba): "several environmental factors, i.e. fisheries interactions, inbreeding, migration, high contaminant loads, higher SSTs [sea-surface temperatures] and limited prey availability may have synergistically interacted to increase the severity of the disease" (p.145-6).Other factors, both natural and anthropogenic, can injure, kill, or cause physiological stress, reduced immune function, and diminished condition in marine mammals, including: anthropogenic noise; severe depletion of food sources resulting in starvation, loss of blubber reserves, and a decline in body condition (e.g. through a mass-mortality of prey or a climate event such an El Niño); and harassment or disturbance (e.g. from fishery or tourism interactions)(Curry 1999; Fair and Becker 2000;Frohoff 2000;St. Aubin 2002; Bejder et al. 2006a,b;Clark et al. 2006). ...
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This technical report reviews findings from an investigation into the mortalities of six bottlenose dolphins (Tursiops aduncus) in the Swan Canning Riverpark in 2009. The report: (a) describes the epidemiology and pathology of these mortalities; (b) presents background information on the ecology of dolphins in the Swan Canning Riverpark and factors known to affect dolphin health; and (c) discusses the potential role of chemical contaminants in the mortalities. These mortalities were investigated in context of dolphin deaths in the Swan Canning Riverpark prior to 2009 and a series of mortalities of dolphins in the Bunbury area between 2008-10, as well as marine mammal mortality events in other locations.
Dolphin: The Myth and the Mammal. H gh -Miff i
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A e, A. 1955. Dolphin: The Myth and the Mammal. H gh -Miff i, B , Ma ach e.
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Gi ch i, 1967 (T, Cha ie );