Ecosys em Se ices 54 (2022) 101406
A ailable online 3 Feb ua y 2022
2212-0416/© 2022 The Au ho s. Published by Else ie B.V. This is an open access a icle unde he CC BY-NC-ND license (h p://c ea i ecommons.o g/licenses/by-
nc-nd/4.0/).
Re iew Pape
Pas , p esen and u u e o he ecosys em se ices p o ided by
ce acean ca casses
M.-Ma ina Quaggio o
a
,
*
, Jos´
e A. S´
anchez-Zapa a
b
, Da id M. Bailey
c
, Ana Payo-Payo
d
,
Joan Na a o
e
, And ew B ownlow
c
, Rob Dea ille
, Se gio A. Lambe ucci
g
, Nu ia Sel a
h
,
Aina a Co ´
es-A izanda
i
,
j
, Fe nando Hi aldo
i
, Jos´
e A. Don´
aza
i
, Ma cos Mole´
on
k
a
Biological and En i onmen al Sciences, Uni e si y o S i ling, S i ling, UK
b
Depa men o Applied Biology, Uni e si y Miguel He n´
andez, Elche, Alican e, Spain
c
Ins i u e o Biodi e si y, Animal Heal h and Compa a i e Medicine, College o Medical, Ve e ina y and Li e Sciences, Uni e si y o Glasgow, Glasgow, UK
d
School o Biological Sciences Zoology Building, Tillyd one A e, Abe deen, Sco land
e
Ins i u de Ci`
encies del Ma (ICM-CSIC), Ba celona, Spain
Ins i u e o Zoology, Zoological Socie y o London, London, UK
g
G upo de In es igaciones en Biología de la Conse aci´
on, INIBIOMA-CONICET, Uni e sidad Nacional del Comahue, Ba iloche, A gen ina
h
Ins i u e o Na u e Conse a ion, Polish Academy o Sciences, K akow, Poland
i
Depa men o Conse a ion Biology, Do˜
nana Biological S a ion (EBD-CSIC), Se ille, Spain
j
Applied Func ional Ecology G oup, Depa men o Plan Biology and Ecology Facul y o Biology, A enida de la Reina Me cedes, 6, 41012 Se ille, Spain
k
Depa men o Zoology, Uni e si y o G anada, G anada, Spain
ARTICLE INFO
Keywo ds:
Ca ion
Dolphin
Ecosys em unc ion
Managemen
S anding
Whale
ABSTRACT
Ecosys em se ices associa ed wi h ce acean s andings ha e been al e ed by humans h ough exploi a ion o
wild popula ions du ing he whaling e a and mo e ecen ly by egula ions on ca cass managemen and disposal
o abide by en i onmen al heal h equi emen s. He e, we sys ema ically e iew he scien i ic li e a u e and
ga he da a on ce acean s andings wo ldwide o: 1) iden i y he ecosys em se ices p o ided by s anded
ce acean ca casses in he pas and p esen ; 2) es ima e he densi y o ce acean s andings cu en ly occu ing in
selec ed coas al a eas a ound he globe, and analyse i s associa ion wi h human popula ion densi y and egu-
la ions; and 3) iden i y and discuss he egula ions and me hods conce ned wi h whale ca cass disposal in speci ic
egions o he wo ld. Ou li e a u e e iew e ealed ha s anded ce acean ca casses ha e p o ided a ich and
a ied a ay o p o isioning, egula ing, cul u al, and suppo ing ecosys em se ices o ancien and mode n
ci ilisa ions wo ldwide. Also, we ound ha he cu en densi y o s anded ca casses (mean: 0.090 s andings •
yea
−1
•km
−1
; ange: 0.001–0.978) and he disposal me hods widely a ied ac oss he s udied egions and
coun ies. In addi ion, nei he human popula ion densi y no he exis ence o egula ions we e good p edic o s o
s anding densi ies. Finally, we p o ide ecommenda ions o he u u e managemen o s anded ce acean
ca casses, by iden i ying hose disposal me hods ha minimize cos s and maximize ecosys em unc ions and
se ices. In pa icula , we encou age na u al decomposi ion in si u whene e possible; o he wise, he p esen
coas al managemen s a egies could be imp o ed by including zoning, seasonal use limi a ion and educa ional
ou each depending upon he local scena io. O e all, u he socio-ecological esea ch is s ongly needed o
guide s anded ce acean ca cass managemen owa ds enhancing he ne bene i s ha humans and ecosys ems
gain om ca casses, especially conside ing ha coas al a eas become mo e popula ed, new disposal egula ions
a e app o ed, and ce acean popula ions a e eco e ing – and hus s andings may become mo e equen .
1. In oduc ion
Mode n socie ies a e inc easingly disconnec ed om na u e
(Dijks a, 2016). One o he many mani es a ions o such disconnec ion
is he g owing implemen a ion o “asep ic” s a egies in na u al esou ce
managemen . Fo ins ance, ca casses o bo h domes ic (Don´
aza e al.,
* Co esponding au ho a : Co ell Building, Biological and En i onmen al Sciences, Facul y o Na u al Sciences, Uni e si y o S i ling S i ling, FK9 4LA, UK.
E-mail add ess: [email p o ec ed] (M.-M. Quaggio o).
Con en s lis s a ailable a ScienceDi ec
Ecosys em Se ices
jou nal homepage: www.else ie .com/loca e/ecose
h ps://doi.o g/10.1016/j.ecose .2022.101406
Recei ed 9 Feb ua y 2021; Recei ed in e ised o m 27 Decembe 2021; Accep ed 12 Janua y 2022
Ecosys em Se ices 54 (2022) 101406
2
2009) and wild animals (Ma galida and Mole´
on, 2016) a e equen ly
emo ed om he ield o minimize disease ansmission isk in many
egions, o en manda ed by egional, na ional o in e na ional egula-
ions. This managemen scena io la gely igno es he pi o al ecological
ole o ca ion (DeVaul e al., 2003; Wilson and Wolko ich, 2011) and
he many con ibu ions ha dead animals can o e o humans (Mole´
on
e al., 2014).
Ce acean ca casses a e a na u ally occu ing elemen o he coas and
humans ha e been exploi ing hem o millennia (See sholm e al.,
2016), p o iding coas al inhabi an s wi h goods such as ood, oil and
bones, bu also wi h knowledge o he sea while obse ing hem
s anded on he sho e (Ellis, 1991). Also, s anded ce aceans a e a key
sou ce o ood o sca enge s, and nu ien s o he sedimen s (e.g. Laid e
e al., 2018; Tucke e al., 2019a), hus being an in eg al pa o coas al
ecosys ems (Panels 1 and 2, Figs. 1 and 2). Howe e , abou o y pe cen
o he human popula ion cu en ly li es wi hin 100 km o he coas , and
as a esul , many coas al a eas ha e been ans o med (C ossland e al.,
2005). The changes include no only he in ense u baniza ion o he
coas and subsequen al e a ion o ecological p ocesses (e.g. Huijbe s
e al., 2013), bu also he applica ion o policies u ging he emo al o
ca casses om he coas al a eas o p o ec human heal h (Tucke e al.,
2018). Combined wi h he pas deple ion o ce acean popula ions
(Hacquebo d, 2001; Panel 3), he exis ing egula ions on emo al o
s anded ca casses has led o adical changes in he abundance and
a ailabili y o la ge ma ine biomass inpu s, which has g ea ly al e ed he
ecosys em unc ions and se ices associa ed wi h ma ine ca ion o e
ime.
In his s udy, we e iew he scien i ic li e a u e and ga he da a on
ce acean s andings o: 1) iden i y he ecosys em se ices p o ided by
s anded ce acean ca casses. He e, ecosys em se ices a e de ined as he
bene i s people ob ain om ecosys ems (Millennium Ecosys em
Assessmen , 2005). These include p o isioning, egula ing, and cul u al
se ices ha di ec ly in luence people, as well as suppo ing se ices (o
ecosys em unc ions) necessa y o main ain he o he se ices. While he
ecen ly de eloped Na u e’s Con ibu ions o People amewo k (NCP)
includes bo h he bene icial (i.e. ecosys em se ices) and de imen al (i.
e. disse ices, con lic s o damages) con ibu ions o na u e o socie ies’
quali y o li e (see Díaz e al., 2018 o an o e iew o he e olu ion o
he concep ), he e we delibe a ely ocus on he bene icial side o
s anded ce acean ca casses; 2) es ima e he densi y o ce acean
s andings occu ing in selec ed coas al a eas a ound he wo ld and
analyse whe he human popula ion densi y and he p esence o egu-
la ions a ec hese eco dings, in o de o gain unde s anding on he
spa ial dis ibu ion and he ac o s a ec ing he po en ial p o ision o
ecosys em se ices by s anded ce aceans; and 3) enume a e he egu-
la ions and me hods conce ned wi h whale ca cass disposal in speci ic
egions o he wo ld, and discuss he p os and cons associa ed wi h each
me hod in e ms o ecosys em se ice p o ision. Based on he indings o
he pape we p o ide ecommenda ions o u u e esea ch and he
Fig. 1. Ce acean ca casses suppo a wide ange o sca enge s, bo h e eb a es
and in e eb a es, in di e en habi a s. See pho o c edi s in Appendix A.
Fig. 2. Andean condo s (Vul u g yphus) used o in ensi ely exploi s anded
whale ca casses. Howe e , he cu en sca ci y o s anded whales has induced
impo an changes in condo s’ beha iou (see Panel 2). Adul Andean condo
soa ing abo e a li ing whale in Pe u (a); se e al Andean condo s o di e en
age eeding upon a s anded whale ca cass in Pa agonia, A gen ina (b). See
pho o c edi s in Appendix A.
M.-M. Quaggio o e al.
Ecosys em Se ices 54 (2022) 101406
3
managemen o ce acean s andings in an inc easingly asep ically
managed wo ld.
2. Ma e ials and me hods
2.1. Li e a u e e iew
We conduc ed a sys ema ic li e a u e e iew acco ding o he
PSALSAR me hod (Mengis e al., 2020), which is an ex ension o he
SALSA amewo k ha has equen ly been used in sys ema ic e iews
(G an and Boo h, 2009) and ha gua an ees concep ualiza ion, accu-
acy and ep oducibili y. In he i s s ep, we de ined ou s udy scope:
ecosys em se ices p o ided by s anded ce acean ca casses. Secondly,
we de ined ou sea ch s ing (see Table B1) and used Web o Science o
sea ch o a icles (published un il 6 h Augus 2020) con aining hese
keywo ds in he en i e a icle. In summa y, ou sea ch p o ocol used he
e ms “humans” and “ecosys em se ices” in combina ion wi h a ious
synonyms o ce acean ca casses (i.e. “s anded whale”, “whale ca cass”,
“s anded ce acean”). This yielded 484 a icles (Table B1), including
epea ed a icles esul ing om di e en sea ches. Thi dly, we es ic ed
he sea ch o pee - e iewed a icles. Then, we sc eened i les and ab-
s ac s o iden i y hose a icles ha deal wi h he ecosys em se ices
p o ided by ce acean ca casses, and we ully ead he selec ed a icles.
This led o a inal selec ion o 27 a icles dealing wi h se ices p o ided
by ce acean ca casses (Table B1). Fou hly, we ex ac ed he ollowing
in o ma ion om he selec ed a icles: 1) ecosys em se ices men ioned
(sepa a ing hem in o p o isioning, egula ing, cul u al and suppo ing
se ices), 2) ce acean species o g oup o which he ecosys em se ices
we e ela ed, 3) s udy a ea, and 4) s udy pe iod (p ehis o ic, his o ic –i.
e. be o e he s a o he 19 h cen u y when he onse o comme cial
whaling occu ed– and mode n –i.e., a e he s a o 19 h cen u y–).
Fi hly, we ca ego ised he selec ed a icles acco ding o hese o me
ac o s (ecosys em se ice ype, ce acean g oup, a ea and pe iod). Spe-
cies we e g ouped acco ding o amilies and pa o de s o Ce acea: a)
baleen whales, b) oo hed whales (excluding small- oo hed whales), and
c) dolphins and po poises, which we e sepa a ed om he o he oo hed
whales because o hei smalle body size and gene ally highe
abundance.
2.2. Da a collec ion on s andings and egula ions
We e ie ed da a on ce acean s anding e en s om 29 coun ies o
egions (Table B3) and egula ions om 23 o hem (Table B4), om an
ini ial lis o 67 coas al coun ies in all con inen s ha we e conside ed
and asked o in o ma ion. Da a on s andings and egula ions we e
di ec ly sha ed ei he by he ma ine mammal ne wo k es ablished in
hese coun ies o by scien is s wo king on ce acean s andings in
di e en academic o ganisa ions, go e nmen al ins i u ions o inde-
penden socie ies. In addi ion, we ex ac ed complemen a y in o ma ion
om local g ey and p ima y scien i ic li e a u e. We selec ed s anding
da ase s ha we e ep esen a i e o he species occu ing in each s udy
a ea and co e ed mul iple yea s. Reco ds o ce aceans ha s anded as
ca casses, died a e s anding o we e unsuccess ully e- loa ed ( e-
s anded and died) we e g ouped as men ioned abo e: a) baleen whales,
b) oo hed whales (excluding small- oo hed whales), c) dolphins and
po poises, and d) uniden i ied ce aceans. Fo each g oup, he o al
numbe o s andings was di ided by he numbe o yea s co e ed by he
da ase and he coas line leng h (km) o he e i o y o coun y (see
mo e de ails in Table B3). Few da ase s also p o ided in o ma ion
ela ed o he a e o he s anded ca casses in e ms o disposal, which
will be summa ised in he Resul s.
2.3. Analysis o s anding da a
To unde s and i s anding epo ing a e in each egion could be
explained by he popula ion densi y and exis ence o s anding
managemen policy o egula ions, we used he 23 egions o which we
we e able o ga he in o ma ion on he exis ence o managemen policy
o egula ions (Table B4), coas leng h and human popula ion densi y.
The exis ence o s anding managemen policy o egula ions was coded
as: “yes”, i p esen and known; “no”, i no p esen and known; and “na”
o unknown ( i e egions). We used he decimal loga i hm o human
popula ion by coas al km
2
as a p oxy o human coas al popula ion
densi y. We pe o med model selec ion using he Akaike’s in o ma ion
c i e ion (AIC; Bu nham and Ande son, 2004). We conside ed models o
be equi alen when he di e ence in AIC wi h he bes model was ΔAIC
<2 (Bu nham and Ande son, 2004). All we e Gene alized Linea Mixed
models (GLMMs) buil in R (R Co e Team, 2013) using a Gaussian link
unc ion.
3. Resul s
3.1. Ecosys em se ices p o ided by ce acean ca casses
Ou li e a u e e iew showed ha ce acean ca casses ha e mainly
supplied p o isioning se ices, ollowed by cul u al, suppo ing and
egula ing se ices (Fig. 3; Table B2). Mos a icles e e ed o mode n
imes, hough hose men ioning p o isioning se ices mos ly ela ed o
p ehis o ic imes (Fig. 3). In ou sea ch, he keywo d “human” was
included in mo e a icles han “ecosys em se ice”. This e eals ha ,
despi e he scien i ic li e a u e acknowledging he associa ion be ween
humans and ce acean s andings, li le o his associa ion is explained in
e ms o ecosys em se ices. He e, we p esen an o e iew o he his-
o ical and cu en ole o ce acean s andings as p o ide s o ecosys em
se ices o humans by compiling he examples p oduced by ou li e -
a u e sea ch and u he pe inen a icles (see also Panel 1).
Ce acean ca casses p o ided ood and aw ma e ial o ea ly ci ili-
sa ions wo ldwide (Fig. 4). Fo ins ance, emains o wo ba nacle spe-
cies, o iginally a ached o he body o sou he n igh whales (Eubalena
aus alis), we e ound in a ca e a a p ehis o ic si e in sou he n Spain,
sugges ing ha p imi i e humans used o consume s anded whale mea
(´
Al a ez-Fe n´
andez e al., 2014). Bone agmen s belonging o ce acean
specimens we e also iden i ied in he Mesoli hic deposi s o o he p e-
his o ic ca es (e.g., G o a dell’Uzzo, I aly; Mannino e al., 2015). The
disco e y, in Angola, o nume ous ools made om whale bone, such as
choppe s and lakes, indica ed ha many componen s o ca casses we e
used in he Lowe Palaeoli hic (Gu ie ez e al., 2001).
In he p e-Columbian e a, indigenous people, such as A awak and
Ca ib g oups om Ba bados (Ca ibbean), also consumed s anded dol-
phins and spe m whales (Physe e mac ocephalus; Rome o and C eswell,
2010). S anded ca casses ha e been o c ucial impo ance o he su -
i al and cul u al de elopmen o o he adi ional coas al communi ies,
such as he Inui li ing in he A c ic egion (Ka ona and Whi ehead,
1988), he “canoe people”, and he Fuegians, inhabi ing he sou he n-
mos Pa agonian coas . The la e conside ed s anded ce aceans a “g ea
Fig. 3. Resul s o he sys ema ic li e a u e e iew on he ecosys em se ices
associa ed wi h ce acean ca casses, acco ding o se ice ype and pe iod o
ime. Ring size is p opo ional o he numbe o a icles ound in he e iew. Fo
mo e de ails, see Table B2.
M.-M. Quaggio o e al.
Ecosys em Se ices 54 (2022) 101406
4
gi o na u e” o be sha ed wi h he en i e communi y (Qui oz e al.,
2016), whe eas Aus alian Abo igines and islande s om he To es
S ai belie ed ha s andings could connec hem o ances al lands,
seas, and hei ances o s (Ha cou e al., 2014). Simila ly, in New
Zealand, M¯
ao i ega ded whale s andings as a symbol o abundance and
ichness because hey we e g ea sou ces o p o ein and ma e ial: hese
esou ces we e ex ac ed ollowing s ic p o ocols o espec hei
spi i ual impo ance (Rodge s, 2017).
In he la e medie al pe iod, bo h Anglo-Saxons and No dic coas al
communi ies used o exploi s anded ma ine mammals (B i o e al.,
2011; Pa sons and Monaghan-B own, 2017). Since he ele en h cen u y,
ce aceans s anded on English sho es ha e been known as “Royal Fish”,
ollowing a s a u e enac ed by Edwa d II (Ga dine , 1997). The ce a-
ceans’ lesh was consumed by weal hy ci izens whe eas he emains o
he ca cass we e ende ed o p oduce oil and he bones used as u ensils
o a is ically ca ed (e.g. he F anks Caske which is a small deco a ed
box made o whale bone om he 8 h cen u y, ep esen ing Anglo-Saxon
a ). Fo he medie al English, whales we e “sea-mons e s” and ac-
co ding o he popula supe s i ion, hei s andings p edic ed he
imminen occu ence o ex ao dina y e en s (Ga dine , 1997; Fig. 4).
As humans de eloped owa ds mode n socie y, hei ela ionship
wi h ma ine mammals also changed. P o isioning ecosys em se ices
a ibu able o s andings we e slowly subs i u ed o cul u al se ices
(Fig. 4). S a ing om he 18 h cen u y, s anded ca casses we e s udied
and La in names we e gi en o ce acean species. S anded ca casses on
beaches, pa icula ly o la ge ce aceans, also equen ly gene a ed high
public in e es (Sousa and B i o, 2012).
F om he 20 h cen u y, i became common o scien is s o collec
s anded specimens o imp o e hei unde s anding o hese aqua ic
mammals (Fig. 4). The skele ons we e collec ed by museums o public
exhibi ion, educa ion (Wei and Pie ce, 2013) o u u e esea ch (Bo -
ella and Bo e o, 2017). Ce aceans and hei ca casses a e now iewed as
indica o s o ocean heal h, and s andings p o ide impo an in o ma-
ion ha enhances ou scien i ic unde s anding o ma ine ecosys ems
and he e ec s o an h opogenic impac s (Bossa , 2011). Recei ing
much a en ion om he media and he public, s andings o la ge
ce acean specimens became sou ces o sus ainable en i onmen al edu-
ca ion. In addi ion, he wo k o olun ee –based o ganisa ions
esponding o ma ine mammal s andings is o g ea alue in aising
public awa eness, which can po en ially enhance animal wel a e and
conse a ion (Roman e al., 2018).
Whale ca casses a e also p o ide s o suppo ing se ices, such as
subsidies o sca enge s. They g ea ly con ibu e o nu ien cycling and
ca bon seques a ion, ep esen ing – in he case o whale alls – a po-
en ial sink o an h opogenic ca bon ha may be impo an o mi i-
ga ing global clima e change (Pe shing e al., 2010; see Panel 1).
3.2. S andings a ound he wo ld
Mean densi y o s anding pe coun y/ egion was 0.09 s andings •
yea
−1
•km
−1
(SD =0.21). No he n Te i o y (Aus alia) had he
lowes annual densi y o s anding e en s (0.001 s andings •yea
−1
•
km
−1
), whe eas Belgium he highes (0.978 s andings •yea
−1
•km
−1
;
Fig. 5). In gene al, as expec ed om hei ela i e abundances (Ham-
mond e al., 2013), dolphins and po poises we e he mos abundan
g oup ac oss all coun ies (Fig. 5), wi h he excep ion o A gen ina and
Chile, whe e baleen whales domina ed he eco ded s anding e en s
(Figu e B.1).
Rega ding he GLMMs explo ing i he s anding densi ies eco ded
could be p edic ed by he p esence o managemen policy o human
popula ion densi y ac oss he s udied a eas, he se o bes compe ing
models (see Table B5) comp ised he model including he e ec o
human densi y (M
1
) and he model including he e ec o legisla ion
p esence (M
2
). Howe e , hese models we e no signi ican ly di e en
om he cons an model, and hei e ec s we e no signi ican (i.e. none
o he be as was signi ican ly di e en om ze o; see Table B5 and
Figu e B.2). Thus, nei he o he abo emen ioned ac o s we e good
p edic o s o eco ded s anding densi ies.
3.3. Regula ions a ound he wo ld
The implemen a ion o egula ions o ca cass managemen is a
Fig. 4. Examples o ecosys em se ices associa ed wi h ce acean ca casses.
P o isioning se ices: p oduc ion o ools (baleen pla es used as b oom; a),
mea (b), e ilize s (c) and oil (oil dish o whale oil; d). Regula ing se ices:
CO
2
seques a ion in he o m o loa ing ca casses be o e sinking (e) be o e
unde going nu ien ecycling h ough ac ion o sca enge s and decompose s
( ). Cul u al se ices: scien i ic ad ancemen in he pas (g) and p esen (h);
p oduc ion o a (F ank Caske , an Anglo-saxon box made o whalebone; i);
educa ional alue in ou doo (j) and indoo museums (m); eco ou ism (k); and
eligious signi icance (l). See pho o c edi s in Appendix A.
M.-M. Quaggio o e al.
Ecosys em Se ices 54 (2022) 101406
5
ecen and ongoing phenomenon. Ou o he 23 s udy coun ies/ egions,
i e did no ha e egula ions ela ed o s andings (22%; Table B4). The
lack o legisla ion was jus i ied by he ins i u ions in ol ed in s anding
managemen because he a eas we e emo e and had low human pop-
ula ion densi y (e.g. New oundland and Lab ado in Canada and
No he n Te i o y). The absence o a s anding esponse ne wo k and a
compe en na ional au ho i y also de e mined he lack o egula ions (e.
g. I eland, which, howe e , has an e icien olun ee ne wo k; IWC,
2016). In some coun ies/ egions, o icial pe missions a e needed o
handle s anded ce aceans in dis ess, o hei ca casses, and speci ic
s akeholde s a e esponsible o he in e en ion and he ca casses’ a e.
Se e al s akeholde s a e in ol ed in hese p ocesses. These may include
local go e nmen s, such as he ci y council and he municipali y, and/o
land manage s, academic ins i u ions, s anding ne wo ks, local NGOs
and museums. Thei ole in he managemen o ca casses a ies om
coun y o coun y. In some coun ies, like New Zealand, indigenous
people a e also in ol ed in he p ocess: o ins ance, M¯
ao i would p ay
o he dead ce aceans be o e hei disposal (Bu e wo h, 2017). P o-
ocols may be implemen ed o guide and acili a e he p ocess o ca cass
disposal (Table B4). This is he case in New Sou h Wales (Aus alia),
whe e he Na ional Pa ks and Wildli e Se ices de eloped a checklis o
guide land manage s when disposing o whale ca casses (NPWS, 2020).
A wide spec um o ca cass disposal me hods exis s, om comple ely
o ela i ely na u al p ocedu es, such as na u al decomposi ion in si u o
he whole ca cass o in pieces, bu ial o he ca cass, owing ou o sea
(o sho e), o land (insho e) o compos ing, o me hods ha in ol e
Fig. 5. Ce acean s anding densi y map (N s andings •yea
−1
•km
−1
) and absolu e abundance o s andings pe g oup o ce aceans (Baleen whales o Mys ice i;
Too hed whales o Odon oce i, excluding small oo hed whales like dolphins and po poises; Dolphins and Po poises; and Uniden i ied ce aceans) o all s udy
coun ies/ egions.
Fig. 6. Di e en me hods o s anded ce acean ca cass disposal and associa ed ecosys em se ices and cos s, o de ed acco ding o hei equency in egula ions o
coun ies/ egions conside ed in his s udy.
M.-M. Quaggio o e al.

Ecosys em Se ices 54 (2022) 101406
6
mo e in e en ion, such as disposal a a local land ill, incine a ion o
ende ing (Fig. 6). In Wes e n Aus alia, mos ca casses a e bu ied
(30%), and only 8% o hem a e le in si u. Excluding he ca casses
whose a e is unknown, in New Zealand and USA mos o he ca casses
a e le on si e o bu ied (NZ: 39% and 38%; USA: 28% and 21%,
espec i ely). Also, in New Zealand, mass s andings ha e o ced au-
ho i ies o use na u al ide d i en decomposi ion. Ca cass ans-
po a ion o land ills o incine a ion plan s is common in many
Eu opean coun ies. Fo ins ance, in Belgium and F ance, which a e
cha ac e ised by high densi ies o s andings, he only op ion is o
des oy ce acean ca casses in app op ia e acili ies. Mul iple disposal
me hods a e allowed in o he Eu opean coun ies (e.g. Spain). In USA,
he hi d mos used me hod is he land ill (18%). Specimens ha will be
used o scien i ic and educa ional pu poses a e ei he collec ed, bu ied
o consumed by sca enge s p io o eco e y o he skele on. In Wes e n
Aus alia and New Zealand, only a small numbe o ca casses a e
collec ed o scien i ic (WA: 0.7%; NZ: 1.3%) o educa ional (WA: 1.2%)
pu poses. When ca casses a e emo ed, hey a e o en nec opsied, so
hey s ill may p o ide a scien i ic and e e ina y se ice. Howe e , in
some cases, he nec opsy pe se is he ul ima e goal, which usually means
mo ing he ca cass o a wildli e escue cen e, whe e i is elimina ed o
cleaned o ob ain he skele on. Nec opsies o la ge whales a e no mally
done in si u.
The me hod o ca cass disposal is con ex -dependen in 45% o
coun ies/ egions, as i may change depending on he exac loca ion
whe e he animal s ands (Table B4). Gene ally, when he ca cass is
ound in a emo e o di icul o access a ea, he common p ocedu e is o
le i decompose na u ally, whe eas i s emo al is conside ed p e e able
when occu ing in a popula ed o ou is ic a ea. This is he case in New
Sou h Wales and Wes e n Aus alia (Aus alia; NPWS, 2020; DPAW).
Howe e , his choice is no con empla ed when he emo al o he
ca cass is compulso y (F ance and Belgium), o he e a e no egula ions
in place because hey a e conside ed unnecessa y (New oundland and
Lab ado , and Falkland Islands). Regula ions a y when s andings
occu in p o ec ed a eas, depending on he po en ial con amina ion and
impac on he exis ing lo a and auna. Special conce ns a ise when
s anded la ge whales a e eu hanized, as oxic chemicals used in some
me hods may pose a possible isk o sca enge s and he su ounding
en i onmen ( elay oxici y), po en ially a ec ing humans nega i ely
(Ha ms e al., 2014). In hese cases, ca casses a e always emo ed o
app op ia e disposal (e.g. UK). In he case o Denma k, emo al o he
ca cass is no loca ion-, bu species-dependen , as only ha bou po -
poises a e le a si e o decompose.
4. Discussion
Ancien and mode n ci ilisa ions wo ldwide ha e bene i ed om he
ich and a ied a ay o p o isioning, egula ing, cul u al, and sup-
po ing ecosys em se ices associa ed wi h s anded ce acean ca casses.
F om Palaeoli hic A icans and Eu opeans o p e-Columbian Ame icans,
and om bo eal Inui s and Icelande s o Paci ic M¯
ao i, p e-indus ial
humans li ing in o nea he coas iewed s andings as a gi om na-
u e (e.g. h al eki in Icelandic, meaning “wind all”, Fielding, 2018), and
ce acean ca casses we e widely used o ood and as sou ce o aw ma-
e ial o hun ing and bu che ing ools and o namen s (e.g. Ga dine ,
1997). Ce aceans also p oduced a deep spi i ual and a is ic inspi a ion
(e.g. Ha cou e al., 2014). Mo e ecen ly, cul u al se ices became he
dominan bene i s associa ed wi h s anded ce aceans (as happened wi h
o he aunal g oups; e.g. Mole´
on e al., 2014), including scien i ic dis-
co e y, in si u and ex si u educa ion, public awa eness aising, and
eco ou ism. O e all, o many ancien and mode n people, ce acean
ca casses signi y an appa en mani es a ion o na u e’s magni icence. All
his e eals he impo an cul u al he i age ha humani y has con-
s uc ed a ound s anded ce aceans. In addi ion, h ough he key
ecological e ec s o ce acean ca casses, which pe ade well beyond he
coas al sys ems and may las o yea s, s andings ha e also led o
ele an suppo ing se ices o pas and p esen ecosys ems. Howe e ,
cu en ly, ecosys em se ices p o ided by s anded ce aceans a e g ea ly
h ea ened by bo h he esul s o educ ion in ce acean popula ions due
o hun ing (see Panel 3) and mode n policies ega ding he disposal o
hese ca casses. F om he In e na ional Whaling Commission 1982
mo a o ium on comme cial whaling, he numbe o coun ies adhe ing
o he In e na ional Con en ion o he Regula ion o Whaling (1946)
inc eased yea a e yea (IWC). In addi ion o his in e na ional
ag eemen , many coun ies ha e pu o wa d na ional legisla ion o
p o ec ma ine mammals, including hei emains. As a esul , he
managemen o any ype o ope a ion in ol ing ma ine mammals ( om
i s in e en ion o disposal o he ca cass) is o en egula ed a na ional
o in a-na ional le els. Because o he na u al ecu ence o s anded
animals on he coas , hese egula ions a e indispensable o no only he
conse a ion o ce aceans and hei ull ecological ole, bu also human
well-being.
Decomposing ce acean ca casses a e a po en ial sa e y haza d o
human heal h, and a nuisance o some locals and beach use s when
occu ing in popula ed a eas (Tucke e al., 2018). Thus, he sys ema ic
emo al o ca casses is o en applied o a oid heal h- ela ed isks and/o
people’s malcon en . The la e could be conside ed he main disse ices
ha s anded ce acean ca casses p o ide o humans. A oiding hese
de imen s, howe e , o en equi es complex logis ics and impo an
economic cos s, which could also be iewed as a so o disse ice. Fo
ins ance, in Aus alia, he beach bu ial o a whale can each up o
app oxima ely $18,000 (USD) when including all he cos s o he ma-
chine y, ope a ions and consul a ion wi h s akeholde s. The anspo
and des uc ion o a small whale a a dedica ed acili y, ins ead, is
a ound $15,000 (USD), wi h cos s a ying acco ding o ca cass size
(Tucke e al., 2018). Mo eo e , as he coas al a eas become “cleaned”
and he coas al ecosys ems “asep ic” o ul il hese human necessi ies,
he s uc u e o na u al communi ies is des abilised and hei ecological
unc ionali y los (Huijbe s e al., 2013), and he p o ision o o he
human demands such as aes he ic enjoymen , educa ion and o he cul-
u al se ices is comp omised. Thus, unde s anding he ecosys em se -
ice po en ial associa ed wi h ce acean s andings is he i s s ep o ully
ecognise and es ablish he di ec and indi ec bene i s o human socie y
and i s needs (Malinauskai e e al., 2021a).
4.1. Policy and managemen ecommenda ions
S anded ce acean ca cass managemen will be an inc easing global
ma e as coas al a eas become mo e popula ed, new egula ions a e
app o ed, and ce acean popula ions eco e (Clapham e al., 1999; see
Panel 3). In addi ion, clima e change and popula ion shi s may po en-
ially lead o s andings in egions which may no ha e p e iously
encoun e ed hem, and s andings may in ol e di e en species han
p e iously seen (Williamson e al., 2021). Thus, managemen s a egies
may need o adap in esponse.
One way o maximise he bene i s de i ed by ce acean ca casses is
iden i ying hose disposal me hods ha can p ese e and/o enhance
ecosys em unc ions and se ices. All he exis ing disposal me hods can
suppo ecosys em se ices, bu some be e han o he s (Fig. 6). Man-
agemen o ce acean ca casses needs o balance hei ecological impo -
ance, he ecosys em se ices hey may p o ide, cos s o managemen
ac ions, biodi e si y conse a ion egula ions and legal and public
heal h equi emen s (Fig. 6). When le in si u, a ce acean ca cass can
p o ide all he ypes o ecosys em se ices, wi h e y low managemen
cos s, bu social discon en may ise. Bu ial is a cos -e ec i e op ion; bu
i he ca cass is bu ied close o he sho eline, he nu ien en ichmen o
he su ounding en i onmen may con amina e he g ound wa e s and
po en ially a ac sha ks o he sho e, which may diminish he ec ea-
ional use o he beach (Tucke e al., 2018; Tucke e al., 2019a; Heiss,
2020). Re u ning he ca cass o he sea by owing i would allow i s
na u al decomposi ion and p o ision o egula ing and suppo ing se -
ices. Howe e , a loa ing ca cass could also become a sa e y haza d in
M.-M. Quaggio o e al.
Ecosys em Se ices 54 (2022) 101406
7
a eas o high ma i ime a ic (Tucke e al., 2018). The ull emo al o
s anded ce aceans om he coas al a ea leads o he loss o ecosys em
se ices and inc ease o he cos s ela ed o hei anspo and disposal.
In pa icula , incine a ion is mo e expensi e han o he me hods and is a
sou ce o g eenhouse gas emissions (Mo ales-Reyes e al., 2015). Among
he op ions equi ing he emo al and anspo o ca casses, compos -
ing cos s (pe uni weigh ) hal o he p ice o he disposal in a land ill o
incine a ion plan (King e al., 2018). Mo eo e , his op ion comes wi h
en i onmen al bene i s oo, e.g. imp o ed soil quali y and b eak-down
o con aminan s and pollu an s (King e al., 2018).
Gi en he p ominen impo ance o s anded ce acean ca casses o
endange ed sca enge s in some a eas/ egions (Chambe lain e al., 2005;
Lambe ucci e al., 2018, see Panel 2) and ecosys em unc ions (Smi h
and Baco, 2003; Bui, 2009; Smi h e al., 2015), we encou age na u al
decomposi ion in si u, p o ided ha legal and sa e y needs a e ul illed.
This measu e, which is he cheapes one, can ob iously be easily applied
in emo e coas s and/o p o ec ed a eas, as some coun ies al eady do. In
mo e popula ed and ou is ic a eas lea ing ca casses in si u may aise
p oblems o sigh , smell, changes in sedimen and g oundwa e chem-
is y, and sha k and e al dog a ac ion (Tucke e al., 2018). In hese
cases, managemen may be season- and/o loca ion-dependen . When
s andings ake place du ing he low- ou ism season, ca casses may be
le in si u. When s andings occu in he high- ou ism season, a empo al
es ic ion, co e ing he main decomposi ion pe iod, could be applied o
he icini y o he ca cass. The implemen a ion o zona ion maps iden-
i ying emo e and u ban beaches could also be implemen ed in o de o
acili a e he decision making. Al e na i ely, ca casses could be chopped
in o smalle pa s o accele a e hei consump ion o decomposi ion.
Ou analysis o s andings a ound he wo ld e ealed ha no all
coun ies o egions will ace he same challenges, as bo h he densi y
and size o he s anded ce aceans we e highly spa ially he e ogeneous.
In gene al, hose coun ies wi h he highes densi ies o s anded ce a-
ceans and mo e es ic i e egula ions o ca cass emo al (e.g. F ance,
Belgium and he Ne he lands) mos ly ecei ed small-sized ce aceans. A
sensible policy migh ake in o accoun bo h he size and numbe o
ca casses. The disposal and anspo o smalle ca casses gene ally cos
less, bo h in e ms o manpowe , logis ics and o e all inancial cos . In a
coun y whe e s andings a e domina ed by small ce aceans such as
dolphins and po poises, a p opo ion o ca casses may, he e o e, be
chosen o be le in si u, o anspo ed o a mo e emo e a ea wi h
limi ed cos s. I bo h small and la ge whales occu a a simila ex en ,
hen logis ics may be eased and cos s educed by emo ing he small size
ca casses, whe eas la ge ca casses could be chopped in si u a e nec-
opsy a oiding anspo cos s, bu po en ially inc easing s a e o s.
Finally, we ecommend ha he pas and p esen alue o s andings,
and ce acean ca casses in gene al, a e included in educa ional ini ia-
i es. In emo e a eas, like in he A c ic egions, in o ma ion on whale
ca casses can be inco po a ed as an educa ional esou ce o he locals
and used o p omo e he inc easing ou is indus y alongside wi h he
one exis ing on li ing whales (Ge aci and Lounsbu y, 1993; Mali-
nauskai e e al., 2021a). This, and o he eco ou ism expe iences, could
be enhanced and expo ed o o he places wo ldwide, as sca enge s
p o ide impo an ec ea ional, aes he ic, lea ning and inspi a ion ex-
pe iences o people (Aguile a-Alcal´
a e al., 2020). An inc eased public
awa eness o he ecological and cul u al ole o ce acean s andings
could help educe nega i e a i udes owa ds hese phenomena and hus
balancing ecosys em se ices and disse ices.
5. Conclusions and u u e esea ch
We ha e p o ided a comp ehensi e o e iew ha could be used o
aise public awa eness o he bene i s ha ce acean ca casses ha e
p o ided o ou ances o s, and he se ices om which mode n humans
could bene i . Thus, u u e s eps should iden i y he main social ac o s
in ol ed and e alua e he public pe cep ion o s andings and associa ed
se ices, which could e en ually imp o e cu en egula ions on ca cass
disposal (Malinauskai e e al., 2021b). Fu he , we encou age he
implemen a ion o inno a i e science ou each ia ci izen science and
educa ion o os e enhanced local ecological knowledge and app ecia-
ion o ecosys em se ices p o ided by ce acean ca casses, which may
lead owa ds mo e sus ainable p ac ices. Finally, bo h use and non-use
alues o ce acean ca casses should be es ima ed in di e en local so-
cial con ex s by in eg a ing ma ke and non-mone a y alua ions o each
ecosys em se ice p o ided (Cook e al., 2020).
We ecommend enhancing he p esen coas al managemen s a e-
gies by widening hei ecosys em pe spec i e by including zoning, sea-
sonal use limi a ion and science ou each. Undoub edly, hese
managemen ac ions need o be suppo ed by a be e scien i ic knowl-
edge o ce acean s andings – and popula ions. Long- e m moni o ing o
s anding e en s, especially i coo dina ed a he sup ana ional le el,
should be a basic componen o he conse a ion o ma ine mega auna
and he ecosys em se ices hey p o ide. These p og ams may supply
de ailed da a on ce acean s anding numbe s and hei spa io empo al
dis ibu ion, which is ele an o e ine ou unde s anding o he ac o s
a ec ing s anding dynamics and o imp o e he e iciency o ca cass
de ec ion and decision making. These da a and knowledge a e also
essen ial o cla i ying he ela ionship be ween s anding e en s and
mul iple explana o y ac o s in addi ion o human densi y and p esence
o egula ions he e explo ed. Impo an ly, his in o ma ion should be
made easily accessible o esea che s. In addi ion, u he e alua ion o
he ecological impac o he di e en managemen echniques o
s andings and compa ing hei cos s and bene i s is essen ial (Tucke
e al., 2018). Only h ough a deepe scien i ic knowledge o he socio-
ecological con ex ha cha ac e izes he ca casses o s anded ce a-
ceans, we will be able o delinea e he mos e icien managemen
s a egies in coas al a eas in an inc easingly popula ed (by bo h humans
and ce aceans) wo ld.
Panel 1. Ecological e ec s o ce acean ca casses.
Whale ca casses a e a signi ican sou ce o ca ion in he ma ine
sys em (Baco and Smi h, 2003), ep esen ing a huge ood supply o
sca enge s and decompose s, bo h ma ine and e es ial (Fig. 1).
“Whale alls” a ac and sus ain a di e se and abundan deep-sea
communi y o e eb a e and in e eb a e sca enge s. By p o iding
in e media e habi a s o be colonised, whale ca casses link chemosyn-
he ic communi ies oge he , enhance he s uc u al complexi y o he
seabed o yea s (Smi h and Baco, 2003) and p omo e adap i e adia ion
o ce ain o ganisms such as he bone-ea ing wo m (Osedax spp.; Smi h
e al., 2015). Mos dead whales sink o he bo om, bu abou 10% may
become buoyan because o gas p oduced du ing decomposi ion
(Pyenson, 2010). While loa ing, ca casses become an easy and ene gy-
ich sou ce o ood o o cas and sha ks (Whi ehead and Ree es, 2005;
Lecle c e al., 2011; Tucke e al., 2019a). O hese, some ca casses
e en ually become s anded on he sho e (Roman e al., 2014). The
ecological impac o hese s anded ca casses on he coas al sca enging
communi y is subs an ial, a ac ing a wide ange o ma ine and
e es ial sca enge s and elici ing nume ical and/o unc ional
esponse in consume s (Polis e al., 1996; Fig. 1). Gian pe els (Wa ham
1990), skuas (L¨
onnbe g 1906) and pola bea s all sca enge on s anded
ce aceans, p o iding he a and p o ein necessa y o cope wi h ex en-
si e pe iods o as ing (Laid e e al., 2018). Whale ca casses may ha e
suppo ed pola bea s in pas wa me in e glacial pe iods, when seal
a ailabili y was limi ed (Laid e e al., 2018), and hus, may play a
c ucial ole in hei adap a ion o ongoing clima e change. A mo e
empe a e la i udes, e es ial sca enge s such as b own bea s, wol es
(Lewis and La e y, 2014), oxes (Ka ona and Whi ehead 1988), hyenas
and jackals (Skinne e al., 1995) also mo e o he coas o eed on dead
ma ine mammals. The ex inc Cali o nia g izzly bea is epo ed o ha e
ed upon salmonids and s anded ma ine mammals (Laid e e al., 2018).
In addi ion, condo s, which a e obliga e sca enge s, la gely specialised
on s anded whales (Chambe lain e al., 2005; Lambe ucci e al., 2018;
o mo e de ails, see Panel 2, Fig. 2). Finally, simila ly o whale alls,
s anding e en s a e likely o en ich subs a a wi h nu ien s (as happens
M.-M. Quaggio o e al.
Ecosys em Se ices 54 (2022) 101406
8
wi h bu ied ce aceans; Bui, 2009) and be discha ged in o coas al wa e s
po en ially a ac ing sha ks (Tucke e al., 2019b; Heiss, 2020). These
nu ien s may also p o ide supplemen s o algae and plan s in he su -
ounding en i onmen , as well as o inc ease landscape he e ogenei y.
Panel 2. Ecological and conse a ion consequences o ma ine
ca cass deple ion: he case o condo s.
Ce acean s andings p o ide long-las ing pulses o ood, which may
de e mine local changes in dis ibu ion and abundance o sca enging
o ganisms and popula ions, as well as changes in he ene gy lux o ood
webs. Whales in pa icula we e an impo an ca ion sou ce o sca -
enge s in his o ic and p ehis o ic imes. Pleis ocene mega aunal
ex inc ion s ongly impac ed e es ial mammals and, o a lesse ex en ,
ma ine mammal popula ions. A e he la e Pleis ocene e es ial
mega aunal ex inc ion in No h Ame ica, he Cali o nia condo (Gym-
nogyps cali o nianus) became ex inc in he a eas whe e hey ed mos ly
on e es ial animals. The condo ’s dis ibu ion ange was es ic ed o
he Paci ic coas al a ea, whe e hey exploi ed he abundan ca ion e-
sou ces p o ided by s andings (Chambe lain e al., 2005). The se e e
deple ion o ma ine mammals du ing he whaling e a o ced condo s o
shi back o e es ial ood esou ces in he la e 1700s, which has
caused a se ious conse a ion p oblem due o lead poisoning om bul-
le s ound in he e es ial ca casses (Chambe lain e al., 2005).
In Pa agonia, Sou h Ame ica, he Andean condo (Vul u g yphus) has
adi ionally nes ed on he Paci ic slope o he Andes, allowing e icien
exploi a ion o he ce acean ca ion ha was once abundan on he coas
(Lambe ucci e al., 2018; Fig. 2). Those b eeding a eas close o he
Paci ic Ocean a e main ained e en oday, when ew whale ca casses a e
a ailable on he wes e n Pa agonian coas . Thus, condo s a e o ced o
ou inely c oss he Andes o he eas o sca enge on ungula e ca casses
om he Pa agonian s eppe, inc easing he ene gy expendi u e and
po en ial isks ha h ea en his species (Lambe ucci e al., 2018).
Popula ions o bo h condo species dec eased un il almos he
ex inc ion in high isk a eas associa ed wi h human impac s. These
h ea s include sa e na i e wild ood educ ions, and he inc easing use
o an h opogenic ood sou ces (Plaza and Lambe ucci, 2020). In he
pas , be o e he se e e declines in whale popula ions, s andings could
ha e been so common ha coas al ecosys em s uc u e and unc ion may
ha e been di e en om hose known oday, and hei e ec s could
ha e e en pe mea ed inland. Unde his scena io, he his o ic lack o
whale s andings has played an impo an ole in he loss o sa e abun-
dan ood sou ces o condo s, and po en ially many o he sca enge s.
Panel 3. The global decline o ce acean popula ions.
Hun ing o whales, especially baleen whales (subo de Mys ice i)
and he spe m whale (Physe e mac ocephalus; subo de Odono oce i),
has been p ac iced o cen u ies by some ci ilisa ions such as he Jap-
anese and Inui (Shoemake , 2005). Mo e ecen ly, whaling s a ed o be
p ac iced also ac oss di e en popula ions such as he Basques in he
11 h cen u y, No semen in he ea ly medie al pe iod, and Du ch, Ca-
ibbeans and Eu opean colonis s o Ame ica in he 15 h cen u y
(Rome o and C eswell, 2010; Pa sons and Monaghan-B own, 2017).
Whaling began in S alba d in 1611 and las ed abou h ee hund ed
yea s. By a ound 1850, he G eenland igh whale was comple ely
emo ed om ma ine ecosys ems a ound S alba d (Hacquebo d, 2001).
The ad en o mode n, comme cial whaling wi h mo e sophis ica ed
essels and he use o explosi es (la e 1800; Ellis, 1991) pe mi ed
whale s o ca ch la ge and as e ce acean species (Pa sons and
Monaghan-B own, 2017). Wi h he expansion in o An a c ic wa e s and
he es ablishmen o whaling s a ions, whaling eached i s his o ical
peak in he ea ly 20 h cen u y. The lack o sus ainabili y o his indus y,
which d as ically educed whale s ocks a a global scale, led i s o he
signing o he Con en ion o he Regula ion o Whaling (Sep . 24, 1931,
155 L.N.T.S 349) in Gene a o An a c ic whale ishe y in 1931, and
e en ually he In e na ional Whaling Commission (IWC) o ake ac ion
and egula e whale ca ches as ea ly as 1946 (IWC). In 1982, a en-yea
mo a o ium was decided and hen applied in 1986 o pe mi whale
popula ions o eco e , bu a he end o i , he pause in comme cial
whaling was ex ended o inde ini e ime (IWC). Cu en ly he e a e 88
pa icipan membe go e nmen s, as Japan wi hd ew i s membe ship in
2013 and esumed whaling in 2019. In addi ion, No way is s ill hun ing
minke whales (Balaenop e a acu o os a a) o in e nal consump ion and
expo o Japan (Lecle c e al., 2011). Iceland and Fa oe Islands also
con inue o p ac ice whaling. Abo iginal subsis ence whaling is also
cu en ly allowed in G eenland, Chuko ka, Bequia and Alaska, al hough
i s e ec on whale popula ions is conside ed negligible (Clapham e al.,
1999).
Wi h millions o ce aceans killed (Clapham e al., 1999), la ge-scale
comme cial whaling is one o he g ea es episodes o unsus ainable
wildli e exploi a ion in human his o y (Clapham e al., 2008). As a
esul , se e al species we e deple ed almos o ex inc ion, al hough some
sign o eco e y ha e been obse ed in some popula ions in ecen de-
cades (Clapham e al., 1999). Because o he slow popula ion g ow h
a e cha ac e ising hese K-s a egy animals, some whale species
(B. musculus, B. physalus and Eubalena aus alis) will ake mo e han a
cen u y o eco e o hal o hei o iginal p e-exploi a ion numbe s,
al hough o he s may eco e soone (e.g. Megap e a no aeangliae; Tul-
loch e al., 2018). The cascading e ec s o he huge dec ease in whale
popula ions ha e p o oundly al e ed he unc ioning o whole ecosys-
ems (e.g. Hacquebo d, 2001).
Al hough i has no been in es iga ed, i can be assumed ha he
numbe o s anded ce aceans has educed in p opo ion o he dec ease
in li ing whales. The diminished a ailabili y o ce acean ca casses has
almos ce ainly also a ec ed he p o ision o ecosys em se ices and
unc ions. The ca bon seques a ion h ough sinking ca casses, o
ins ance, would be an o de o magni ude g ea e i whale popula ions
we e es o ed o p e-whaling numbe s (Pe shing e al., 2010).
Decla a ion o Compe ing In e es
The au ho s decla e ha hey ha e no known compe ing inancial
in e es s o pe sonal ela ionships ha could ha e appea ed o in luence
he wo k epo ed in his pape .
Acknowledgemen s
This s udy was suppo ed by he Spanish Minis y o Economy, In-
dus y and Compe i i eness and EU ERDF unds h ough he p ojec s
CGL2015-66966-C2-1-2-R and CGL2017-89905-R. M.M. was suppo ed
by a esea ch con ac Ram´
on y Cajal om he MINECO (RYC-2015-
19231), and A.CA. by a con ac Juan de la Cie a Inco po aci´
on (IJCI-
2014-20744; Spanish Minis y o Economy and Compe i i eness, Spain)
and a Pos Doc con ac P og ama Viçen Mu o Go e n Balea , Spain
(PD/039/2017). NS was suppo ed by he Bekke p og amme o he
Polish Na ional Agency o Academic Exchange (NAWA). Acknowl-
edgemen s o speci ic ins i u ions and people sha ing hei aluable
s andings da abase is in Table B3.
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