297
Re is a Chilena de His o ia Na u al
76: 297-312, 2003
The dis ibu ion o li o al cap ellids (C us acea: Amphipoda:
Cap ellidea) along he Paci ic coas o con inen al Chile
La dis ibución de cap élidos li o ales (C us acea: Amphipoda: Cap ellidea) en la
cos a del Pací ico de Chile con inen al
MARTIN THIEL1* , JOSÉ M. GUERRA-GARCÍA2, DOMINGO A. LANCELLOTTI1 & NELSON VÁSQUEZ1
1Facul ad de Ciencias del Ma , Uni e sidad Ca ólica del No e, La ondo 1281, Coquimbo, Chile;
e-mail: [email p o ec ed]
2 Labo a o io de Biología Ma ina, Depa amen o de Fisiología y Biología Animal, Facul ad de Biología,
Uni e sidad de Se illa, Apa ado 1095, E-41080, Se illa, España
(*co esponding au ho )
ABSTRACT
Many li o al cap ellid species ha e a e y ample dis ibu ion, some ha ing been epo ed om all o e he wo ld. The
cosmopoli an dis ibu ion o many li o al cap ellid species migh be acili a ed by he ac ha hey a e o en associa ed
wi h ouling communi ies on loa ing objec s, which ha e a high po en ial o a - ange dispe sal. This dispe sal po en ial
may also ha e implica ions o he dis ibu ion o cap ellids on local and egional scales. He ein we examined he
dis ibu ion o li o al cap ellid species on wo spa ial scales, local ( ens o kilome e s) and egional (hund eds o
kilome e s) along he Paci ic coas o con inen al Chile. On he local scale, we s udied he cap ellid auna in di e en
habi a s (in e idal boulde s, sub idal algal and seag ass beds, ouling communi y on buoys and opes); on he egional
scale we ocused only on cap ellids associa ed wi h he ouling communi y on buoys and opes. We ound a o al o six
cap ellid species, some o which we e e y abundan bo h on he local as well as on he egional scale. On he local scale
we ound a di e ence be ween he h ee s udied habi a ypes wi h espec o he assemblage o cap ellid species, some
o which we e ound in mo e han one habi a . The highes species ichness and abundance o cap ellids was ound in
he ouling communi y associa ed wi h ancho ed buoys and opes. On he egional scale we ound e y high numbe s
o cap ellids in he ouling communi y o he no he n egion (n o 30° S), and dec easing abundances and species
ichness in he cen al egion (30–37° S). No cap ellids we e ound in he sou he n egion o he s udy a ea (37–42° S).
This pa e n coincides wi h he global dis ibu ion o li o al cap ellid species, which a e mos abundan and di e se a
low la i udes bu occu in low abundances and low di e si y a high la i udes. De ached buoys ha we e ound a ew km
o he coas ha bo ed simila cap ellid assemblages (including o ige ous emales) as ancho ed buoys, sugges ing ha
buoys and o he loa ing subs a a (li e , mac oalgae) may acili a e dispe sal o cap ellids (and o he epibio a) along
he Paci ic coas o Chile. Since a i icial and na u al loa ing subs a a a e also abundan a high la i udes i is in e ed
ha he low di e si y o li o al cap ellids a high la i udes is no due o lack o dispe sal ec o s, bu a he o o he
ac o s.
Key wo ds: C us acea, Amphipoda, Cap ellidea, Chile, biogeog aphy.
RESUMEN
Muchas especies de cap élidos del li o al p esen an una amplia dis ibución, algunas han sido ci adas a lo la go de odo
el mundo. La dis ibución cosmopoli a de muchas especies de cap élidos pod ía debe se al hecho de que se asocian
ecuen emen e a las comunidades “ ouling” en obje os lo an es, que p esen an una capacidad impo an e de dispe sión
a la gas dis ancias. Es a capacidad de dispe sión puede ene implicaciones en la dis ibución de cap élidos a escala local
y egional. En es e es udio, noso os examinamos la dis ibución de las especies de cap élidos li o ales en dos escalas
espaciales, local (decenas de kilóme os) y egional (cien os de kilóme os), a lo la go de la cos a Pací ico de Chile
con inen al. A escala local se es udió la auna de cap élidos en dis in os hábi a s (bolones in e ma eales, p ade as de algas
y ane ógamas ma inas, comunidades “ ouling” de boyas y cue das); a escala egional el es udio se cen ó solamen e
en los cap élidos asociados a boyas y cue das. Se encon a on un o al de seis especies de cap élidos, algunos de los cuales
ue on muy abundan es a escala local y egional. A escala local, exis ió una di e encia en e los es ipos de hábi a con
espec o a la auna de cap élidos, algunos de los cuales es u ie on p esen es en más de un hábi a . Los alo es más al os
de iqueza especí ica y abundancia se encon a on en la comunidad “ ouling” asociada a boyas y cue das ancladas. A
escala egional, los cap élidos ue on más abundan es y di e sos en las comunidades “ ouling” de la egión no e (n de
30º S), mien as que la abundancia y iqueza de especies disminuye on en la egión cen al (30-37º S). No se encon a on
cap élidos en la egión su del á ea de es udio (37-42º S). Es e pa ón coincide con la dis ibución global de especies de
cap élidos li o ales, mos ando los alo es más al os de abundancia y iqueza de especies en las bajas la i udes y los
alo es más bajos en las al as la i udes. Boyas desp endidas desde sus ama as y encon adas a a ios km de la cos a
albe ga on asociaciones de cap élidos (incluyendo hemb as o íge as) simila es a las de las boyas ijadas, sugi iendo que
298 THIEL ET AL.
INTRODUCTION
Cap ellid amphipods a e common membe s o
many li o al habi a s, and hey a e pa icula ly
abundan in epibio ic ouling communi ies. They
a e e y common and di e se on e ec b yozoans
and hyd ozoans and on plan subs a a such as
mac oalgae and seag asses (McCain 1968). They
eed on suspended ma e ials, p ey on o he o gan-
isms, o g aze on epibio ic auna and lo a (Caine
1974). Locally, cap ellids a e impo an p ey o
many coas al ish species (Caine 1987, 1989,
1991), and hey may be impo an membe s o
li o al communi ies.
Many cap ellid species ha e a wide dis ibu ion
and he e a e many examples o species ha a e
conside ed o be cosmopoli ans (Takeuchi &
Sawamo o 1998). Wi h hei gene al mo phol-
ogy, cap ellids a e well adap ed o cling o sub-
s a a such as algae and hyd oids (Takeuchi &
Hi ano 1995). Using hei las pe eiopods hey
can i mly hold on o b anches o algae, b yozoans
and hyd ozoans. The pleopods, which a e used
o swimming in o he amphipod c us aceans, a e
educed in cap ellids. The e o e, al hough
cap ellids can swim (Caine 1979), hey p obably
a e no e y e icien swimme s. This, as well as
he lack o a plank onic la al s age, sugges s ha
cosmopoli an cap ellid species may be dis ib-
u ed passi ely by clinging o loa ing ma e ials
a he han by ac i e swimming. Floa ing ma e i-
als such as mac oalgae a e easily dis ibu ed be-
ween dis an loca ions (Ingól sson 1995, Hobday
2000a) and cap ellids and o he amphipods a e
commonly ound on his ype o subs a a (Hobday
2000b). Many cap ellid species a e also e y
unselec i e wi h espec o hei subs a um and
hey colonize a wide a ie y o di e en sub-
s a a. Fo example he species C. equilib a has
been ound on algae, seag asses, b yozoans,
bi al es, sponges and o he subs a a (K app-
Schickel 1993). This low selec i i y o a wide
a ie y o subs a a ha ha e high loa ing po en-
ial sugges s ha hese species migh ha e a wide
dis ibu ion, pa icula ly in egions whe e ocean
cu en s anspo suspended o loa ing ma e i-
als (algae, b yozoans, wood o buoys and o he
an h opogenic ma e ial) o e la ge dis ances.
Al hough he abundance and species ichness
o cap ellids in many a eas o he wo ld’s oceans
las boyas y o os sus a os lo an es pueden acili a la dispe sión de cap élidos (y o a epi auna) a lo la go de la cos a
Pací ico de Chile. Teniendo en cuen a que los sus a os lo an es na u ales y a i iciales son ambién abundan es en al as
la i udes se in ie e que la baja di e sidad de cap élidos li o ales en al as la i udes no se debe a la al a de ec o es
dispe san es sino a o os ac o es.
Palab as cla e: C us acea, Amphipoda, Cap ellidea, Chile, biogeog a ía.
is s ill poo ly known, Laubi z (1970) poin ed ou
ha su ace wa e empe a u e is an impo an
ac o de e mining he dis ibu ion o li o al
cap ellids along he Ame ican coas o he No h
Paci ic. Simila as o o he axa (e.g., co als –
F ase & Cu ie 1996; molluscans – Roy e al.
1998, 2000), species ichness o cap ellids de-
c eases g adually owa ds no he n la i udes
(Laubi z 1970). Wa e empe a u e may a ec he
dis ibu ion o cap ellids along he Ame ican coas
o he Sou h Paci ic in a simila way as in he
No h Paci ic since he oceanic egime and in
pa icula he la i udinal a ia ion in wa e em-
pe a u es show simila i ies be ween hese wo
egions. I can he e o e be expec ed ha di e -
si y o li o al cap ellids along he Paci ic coas
o Chile also dec eases owa ds high la i udes.
Ne e heless, on a mo e local scale, and in pa -
icula in bay sys ems, wa e ci cula ion leads o
in ense local mixing (Soba zo 2002), and he
dis ibu ion pa e n o cap ellid amphipods he e-
o e can be expec ed o be ela i ely uni o m
wi hin local bay sys ems.
In he p esen s udy, we examined he composi-
ion o he cap ellid auna bo h on a local and
egional scale. The local dis ibu ion was exam-
ined wi hin a sys em o bays, i.e., o e a scale o
ens o km, while he egional dis ibu ion o
cap ellids was examined along a la i udinal g a-
dien a si es sepa a ed by hund eds o km. The
p incipal objec i e o his s udy was o e eal he
species ichness o li o al cap ellids and hei
dis ibu ion pa e n along he Paci ic coas o
con inen al Chile. We inco po a e da a on he
cap ellid assemblage om de ached buoys ound
a ew km o sho e since his in o ma ion con ib-
u es o a be e unde s anding o bo h he local
and egional dis ibu ion o cap ellids.
MATERIAL AND METHODS
The local dis ibu ion pa e n o li o al cap ellids
was examined in he bay sys em o Coquimbo,
no he n-cen al Chile (Fig. 1). In No embe /
Decembe 2000, semi-quan i a i e samples we e
aken in h ee di e en habi a s, in e idal boul-
de s, sub idal algal and seag ass beds and ouling
communi ies om buoys and opes. P elimina y
su eys had e ealed ha cap ellid amphipods
299DISTRIBUTION OF LITTORAL CAPRELLIDS FROM CHILE
we e mo e common in hese han in o he li o al
habi a s (Gue a-Ga cía & Thiel 2001). A each
o he se en sampling si es we a emp ed o sample
all h ee habi a s keeping he minimum dis ance
be ween di e en habi a s < 1,500 m; a some
si es no all habi a s we e ep esen ed. Sessile
auna oge he wi h any adhe ing amphipods was
sc aped om boulde s loca ed in he low in e -
idal zone. We ocused on a eas ha we e colo-
nized by hyd ozoans Obelia dicho oma, which we
sc aped om he unde side o se e al boulde s
un il comple ing a olume o 10-20 mL. In he
sub idal beds o G acila ia chilensis and
He e ozos e a asmanica we collec ed he p inci-
pal plan species oge he wi h o he occasional
algae (Callophyllis a iega a, Chond acan hus
chamissoi, Sa codio heca gaudichaudii,
Sphae ococcus co onopi olius and Halop e is
sp.). Plan s we e aken om he bo om by a semi-
au onomous di e and placed in a sampling ja
un il comple ing a olume o app oxima ely 1,000
mL. F om buoys and opes associa ed wi h aquac-
ul u e o ha bo s uc u es, we collec ed he mos
common epibio a (me azoans such as Pyu a
chilensis, Bugula ne i ina and B. labella a,
Tubula ia c ocea, ci ipeds om he amily
Balanidae; and mac oalgae such as Ul a lac uca,
Polysiphonia panicula a and P. mollis,
Chond acan hus chamissoi, Lessonia abecula a,
o en co e ed wi h b yozoans o he genus
Memb anipo a) oge he wi h he adhe ing am-
phipods. Samples we e ei he aken by a semi-
au onomous di e who sepa a ed he ouling o -
ganisms om he buoys and opes and placed
hem in a sampling ja a he wa e su ace. Al e -
na i ely, buoys and opes we e pulled in o a boa
Fig. 1: Bay sys em in es iga ed in he p esen s udy. Samples we e aken om buoys, seag ass and algal
beds, and boulde s on exposed ocky sho es along s a ions A-G; he h ee di e en subs a a we e no
p esen a all sampling s a ions. Do s along he Paci ic coas o con inen al Chile indica e sampling si es
o egional s udy.
Sis ema de bahías in es igado en el p esen e es udio. Las mues as ue on ecolec adas en boyas, p ade as y bolones a lo
la go de las es aciones A-G. Los es subs a os conside ados no siemp e es u ie on p esen es en odos los pun os de
mues eo. Los cí culos a lo la go de la cos a Pací ico de Chile con inen al indican los si ios de mues eo del es udio
egional.
300
whe e ouling o ganisms we e sepa a ed and im-
media ely placed in he ja . Fouling o ganisms
we e ga he ed om di e en buoys and opes
un il comple ing a olume o 500 mL.
The main objec i e o his su ey was o de e -
mine he species ichness o li o al cap ellids in
di e en habi a s. In o de o ob ain ep esen a-
i e es ima es o species, we selec ed di e en
sample olumes in each habi a (20 mL unde
boulde s, 500 mL in buoys and opes and 1,000
mL in algal beds) he eby aking s uc u al di e -
ences among habi a s in o accoun . P elimina y
sampling had indica ed ha unde boulde s
cap ellids cling o small hyd oids o o he sub-
s a um di ec ly, while on algae hey oam o e
la ge a eas o algal su ace. A su ey by Gue a-
Ga cía & Thiel (2001) in he bay sys em o
Coquimbo indica ed ha he selec ed sampling
olumes a e su icien o ob ain ep esen a i e
es ima es o species ichness in he habi a s s ud-
ied he ein. Since his p elimina y su ey had
e ealed la ge di e ences in cap ellid abundance,
he ein we used a semi-quan i a i e me hod in o -
de o es ima e cap ellid abundances in he di e -
en habi a s: p esen (1-10 indi iduals sample-1),
common (11-100 indi iduals sample-1), abundan
(101-1,000 indi iduals sample-1) and supe abun-
dan (> 1,000 indi iduals sample-1). We exp essed
he abundance o cap ellids as indi iduals pe
sample and no as indi iduals pe sample olume
he eby aking in o accoun he di e en habi a
s uc u e. The use ulness o a semi-quan i a i e
scale in mul i a ia e analysis has been demon-
s a ed in ecological s udies dealing wi h ma ine
in e eb a es (Maldonado & U iz 1995, Ca ballo
e al. 1996, Na anjo e al. 1996, Gue a-Ga cía
2001). Ne e heless, acco ding o Cla ke &
Wa wick (1994), when sampling a i ac s ende
quan i a i e compa isons un easible, a educ ion
o simple p esence o absence o each species is
ecommended. Following hese sugges ions, simi-
la i ies o cap ellid occu ence among di e en
habi a s on he local scale we e examined by
means o a classi ica ion analysis based on p es-
ence/absence da a ins ead o he semi-quan i a-
i e da a. The Jacca d simila i y index (Jacca d
1908) and he UPGMA (unweighed pai -g oup
a e age me hod) agg ega ion algo i hm (Snea h
& Sokal 1973) we e used. The signi icance o he
esul ing clus e s was es ablished ollowing he
boo s ap me hodology as in Lancello i & T ucco
(1993) (a e Jaksic & Medel 1987). F om he
ini ial species*habi a ma ix wi h p esence/ab-
sence da a, 1,000 s ochas ic eassignmen s we e
made, compu ing on each occasion simila i y
pseudo- alues o each possible combina ion o
habi a s. All pseudo- alues gene a ed a e 1,000
shu les we e plo ed in a equency his og am,
es ablishing he signi icance, i.e., a a 0.05 o
0.10 α-le el, a he 95 h o 90 h pe cen ile, espec-
i ely. The simila i y alue o which he 95 h
pe cen ile co esponded was conside ed he c i i-
cal alue. Thus, o any clus e , a simila i y alue
abo e his c i ical alue was conside ed highe
han expec ed a andom, and i was conside ed
signi ican a he α = 0.05 le el.
Fo he s udy o he egional dis ibu ion o
cap ellids we ocused on he species associa ed
wi h he ouling communi y om buoys and opes,
since p e ious s udies had e ealed ha his is a
e y impo an habi a o li o al cap ellid spe-
cies (Bynum 1978, Caine 1987). A 12 di e en
si es be ween 18.4° and 41.9° S, spanning ~ 3,000
km along he Paci ic coas o con inen al Chile
(Fig. 1), samples we e aken om buoys and
opes associa ed wi h aquacul u e o ha bo s uc-
u es. Fo Coquimbo we conside ed he esul s
om Bahía La He adu a (si e C in Fig. 1). Sam-
pling was done du ing aus al summe 2001 in he
same way as desc ibed abo e. Fouling o ganisms
we e sepa a ed om di e en buoys and opes
and placed in o he sampling ja un il comple ing
a olume o 500 mL. In his way we ob ained one
cumula i e sample o each o he 12 di e en
si es. All samples we e ixed wi h 5 % o malin
and la e washed o e a mesh o 250 µm be o e
so ing hem in he labo a o y. The B ay-Cu is
dis ance index (B ay & Cu is 1957) and he
UPGMA (unweighed pai -g oup a e age me hod)
agg ega ion algo i hm (Snea h & Sokal 1973)
we e used. The analysis was conduc ed using he
semi-quan i a i e da a, i.e., wi h an o de o mag-
ni ude co esponding o a loga i hmic ans o -
ma ion, which is ypically used o his so o
analysis in o de o s anda dize in e speci ic a i-
abili y (Cla ke & Wa wick 1994). Be o e conduc -
ing he analysis we added he alue o 1 x 10-5 o
each elemen in he o iginal ma ix, which al-
lowed o inco po a e he locali ies wi h no
cap ellids (ze o alues) in o he analysis – his
p ocedu e p oduced no changes in he gene al
associa ion pa e n. The signi icance o he clus-
e s esul ing om his analysis was es ablished
ollowing he same p ocedu e as on he local
scale bu based on he species*si e semi-quan i a-
i e da a ma ix.
Du ing a wo-day oceanog aphic c uise con-
duc ed du ing he aus al all 2002 in coas al
wa e s o he Bay sys em o Coquimbo we col-
lec ed six de ached buoys oge he wi h hei oul-
ing communi y. Addi ionally, du ing se e al
oceanog aphic c uises along he Chilean coas ,
which we e conduc ed du ing aus al summe 2002
(nine c uises be ween A ica, 18° S, and Isla Mad e
THIEL ET AL.
301
de Dios, 50° S, each c uise 10-30 Nm o he
coas ), we collec ed wo de ached buoys o he
coas a app oxima ely 27° S. On mos occasions
we sampled he en i e buoy and a e washing i
o e a mesh o 250 µm we sc aped he ouling
communi y om he buoys. Due o logis ic ea-
sons di e en olumes we e sampled o he oul-
ing communi y om hese de ached buoys. In all
cases he semi-quan i a i e es ima es ep esen
minimum es ima es o he o al cap ellid abun-
dance on each buoy. Following p ese a ion in
he ield, hese samples we e p ocessed in he
labo a o y in he same way as desc ibed abo e.
De ailed in o ma ion abou he de ached buoys
can be ound in Appendix 1.
RESULTS
Local dis ibu ion o cap ellids om di e en
li o al habi a s
A o al o six cap ellid species we e collec ed in
di e en li o al habi a s along he bay sys em o
Coquimbo. None o hese species was ound in all
sampled habi a s, bu some species (Cap ella
equilib a Say, 1818, C. scau a Temple on, 1836,
Deu ella enenosa Maye , 1890) occu ed in wo
o he h ee s udied habi a s, while o he s
(Cap ellina longicollis (Nicole , 1849), Cap ella
e ucosa Boeck, 1871, Pa acap ella pusilla
Maye , 1890) we e only ound in one o hese
habi a s (Table 1). The species C. scau a was one
o he mos widesp ead species, being common in
almos all samples om buoys and p esen in all
samples om algal and seag ass beds. Al hough
he p esence o C. scau a in algal and seag ass
beds was cons an along he bay sys em s udied,
i s abundance was compa a i ely low, ne e ex-
ceeding 10 indi iduals pe sample. The species
D. enenosa was e y abundan on buoys and did
also occu unde in e idal boulde s. Cap ella
e ucosa also was e y abundan on buoys bu
did no occu in any o he o he wo habi a s.
Cap ella equilib a was p esen on some buoys,
bu also occu ed unde boulde s oge he wi h D.
enenosa, bo h species being associa ed wi h he
hyd oid Obelia dicho oma. The p esence o
Pa acap ella pusilla was es ic ed o he boul-
de s om sampling si e C. Cap ellina longicollis
TABLE 1
Abundance o cap ellid species in he di e en habi a s (1 - buoys, 2 - algal and seag ass
beds, 3 - boulde s) and he sampling si es (A-G) in he bay sys em o Coquimbo; (*) indica es
he seag ass bed; sample olume o buoys is 500 mL, o algal and seag ass beds is 1,000 mL
and o boulde s app oxima ely 20 mL. Fo u he de ails see ex
Abundancia de las especies de cap élidos en los dis in os hábi a s (1 – boyas, 2 – p ade as, 3 - bolones) y es aciones
mues eadas (A-G) a lo la go del sis ema de bahías de Coquimbo; (*) indica la p ade a de ane ógama. El olumen
de mues eo pa a boyas ue 500 mL, pa a p ade as 1.000 mL y pa a bolones ap oximadamen e 20 mL. Ve ex o pa a
más de alles
Buoys Algal and seag ass beds Boulde s
A1 B1 C1 E1 F1 G1 A2 B2 C2 D2 E2 F2 G2* A3 C3 D3 E3 F3
Cap ellina longicollis
Cap ella equilib a
Cap ella scau a
Cap ella e ucosa
Deu ella enenosa
Pa acap ella pusilla
(A) Bahía de Coquimbo, no h Abundance pe sample:
(B) Bahía de Coquimbo, sou h P - p esen (1-10 indi iduals)
(C) Bahía de La He adu a Co - common (11-100 indi iduals)
(D) Bahía de To o alillo Ab - abundan (101-1,000 indi iduals)
(E) Bahía de Guanaque os
(F) Bahía de Tongoy, no h
(G) Bahía de Tongoy, sou h
(*) Seag ass bed (He e ozos e a asmanica)
DISTRIBUTION OF LITTORAL CAPRELLIDS FROM CHILE

302
was, oge he wi h C. scau a, he only species
eco ded om he algal and seag ass beds.
The epi ouling communi y om buoys and opes
was he habi a wi h he highes species ichness
and abundance o li o al cap ellids. The cap ellid
auna di e ed among he s udied habi a s, bu on
a local scale ( ens o km) li le a ia ion was
ound wi hin habi a s, wi h he excep ion o boul-
de s (Fig. 2). The samples om buoys and algal/
seag ass beds we e g ouped in wo sepa a e
g oups, bo h connec ing wo subg oups. While
he g oup connec ing cap ellid auna om buoys
was signi ican (P < 0.05), hose om he algal/
seag ass beds o med dis inc subg oups a 0.10 >
P > 0.05. In con as , samples om boulde s
o med no dis inc g oup sugges ing a compa a-
i ely he e ogeneous habi a , possibly exposed o
equen ex inc ion/coloniza ion e en s.
Regional dis ibu ion o cap ellids associa ed wi h
ancho ed buoys
Fou cap ellid species we e ound on he an-
cho ed buoys along he Chilean coas (Fig. 3).
Simila as in he bay sys em o Coquimbo,
Cap ella e ucosa and C. scau a we e he mos
abundan species on buoys, being widely dis ib-
u ed in he no he n egion o he Chilean coas .
Cap ella equilib a was only p esen in a ew
locali ies and he species Deu ella enenosa only
occu ed in Coquimbo. Bo h, abundance and spe-
cies ichness o cap ellids on he buoys and opes
dec eased om no h o sou h, and no cap ellids
we e ound in he sou he n egion (a 39.8° and
41.9° S) (Fig. 3). The classi ica ion analysis
showed h ee dis inc clus e s, one in he no he n
egion cha ac e ized by ela i ely high species
ichness and abundances, whe e wo subg oups
a e connec ed wi h P < 0.10, one in he cen al
egion wi h highe he e ogenei y, whe e species
ichness and abundance dec eased, and one in he
sou h whe e no li o al cap ellids we e ound in
he ouling communi y associa ed wi h buoys (Fig.
4).
Some in aspeci ic mo phological a ia ion has
been ound o Cap ella scau a and C. e ucosa
a some si es. The s iking head spine, ypical o
C. scau a epo ed h oughou he wo ld, was
e y educed in he specimens collec ed om Isla
S a. Ma ia (23.4° S). Ne e heless, examina ion
o an ennae, gna hopods, mou hpa s and abdo-
men o hese specimens con i med ha hey be-
longed o C. scau a. In he specimens o Cap ella
e ucosa om Cale a Tumbes (36.6° S), he
sou he nmos s a ion whe e cap ellids we e ound,
he ube cles on he do sal pe eoni es we e ab-
sen . Taking in conside a ion he ea u es o an-
ennae, hese specimens we e assigned o C.
e ucosa. These obse a ions, howe e , con i m
he g ea need o u u e axonomic s udies in he
complex o species simila o C. e ucosa (see
also Gue a-Ga cía & Thiel 2001). Rega dless o
he mo phological a ia ions in some o he speci-
mens ound he ein, a s ong dec ease in abun-
dance and species ichness o li o al cap ellid
owa ds highe la i udes was ound (Fig. 3).
Cap ellids associa ed wi h de ached (= pelagic)
buoys
On he six de ached buoys ha we e collec ed
while loa ing in coas al wa e s o he Bay sys-
em o Coquimbo, a o al o ou di e en cap ellid
species was ound (Table 2). The mos common
cap ellid species we e Cap ella scau a and C.
Fig. 2: Clus e analysis showing he ela ionships
among samples on he local scale. Simila i ies
based on p esence/absence o cap ellid species on
di e en subs a a (buoys: 1, algal/seag ass beds:
2; boulde s: 3) and sampling s a ions (A-G).
Dashed lines indica e signi ican le els: G oups o
habi a s 1 (buoys) and 2 (algal/seag ass beds)
signi ican a 0.10 > P > 0.05.
Análisis de conglome ados mos ando las elaciones en e
subs a os en una escala local. Las simili udes es án
basadas sob e la p esencia/ausencia de cap élidos en los
di e en es subs a os (boyas: 1, p ade as: 2; bolones: 3) y
es aciones de mues eo (A-G). Las líneas somb eadas
indican ni eles de signi icancia: los g upos de hábi a s 1
(boyas) y 2 (p ade as) ue on signi ica i os a una p obabili-
dad 0,10 > P > 0,05.
THIEL ET AL.
303
equilib a, which we e each ound on ou o he
six buoys. The o he wo cap ellid species, C.
e ucosa and Deu ella enenosa, we e each ound
on wo buoys. Abundances o indi idual cap ellid
species equen ly exceeded 100s o indi iduals
pe buoy (Table 2). On all buoys, we also ound
o ige ous emales.
Du ing he nine esea ch c uises be ween A ica
and Isla Mad e de Dios we ound wo de ached
buoys loa ing in coas al wa e s o Calde a. Each
o hese wo buoys ha bo ed h ee cap ellid spe-
cies (C. equilib a, C. scau a and C. e ucosa)
(Table 2). Abundances o C. equilib a was high-
es on he buoys i sel , while C. scau a domina ed
on he ope s ill a ached o one o hese wo
buoys. O all h ee cap ellid species and on each
o he wo buoys, o ige ous emales we e ound.
DISCUSSION
The esul s o he p esen s udy demons a ed ha
he cap ellid auna along he Paci ic coas o
con inen al Chile is compa a i ely poo . Despi e
a ela i ely la ge sampling e o on he local and
egional scale only six species o li o al cap ellids
we e ound. Rega dless o he low species num-
be , dis inc geog aphic pa e ns we e ound. In
Fig. 3: Abundance and dis ibu ion o he cap ellid species associa ed wi h he ouling communi y o
buoys and opes along he Paci ic coas o con inen al Chile. A each si e, epi aunal o ganisms om
di e en buoys and opes we e ga he ed un il comple ing a olume o 500 mL.
Abundancia y dis ibución de los cap élidos asociados a la comunidad epi i a de boyas y cue das a lo la go de la cos a
Pací ico de Chile con inen al. En cada localidad, se ecolec a on o ganismos epí i os de di e en es boyas y cue das has a
comple a un olumen de 500 mL.
DISTRIBUTION OF LITTORAL CAPRELLIDS FROM CHILE
304
he ollowing we will compa e he dis ibu ion
pa e ns o li o al cap ellids on di e en geo-
g aphic scales, and we discuss he ac o s in lu-
encing hese pa e ns.
Local dis ibu ion o cap ellid species in he bay
sys em o Coquimbo
Wi hin he bay sys em examined he ein, mos
cap ellid species we e ound in wo ypes o habi-
a . Those habi a s we e e y di e en (e.g., oul-
Fig. 4:Clus e analysis showing he ela ionships among sampling si es along he Paci ic coas o
con inen al Chile ( egional scale). Dis ance was based on semi-quan i a i e alues o cap ellid
abundance. Dashed lines indica e signi ican le els: g oup o sampling si es om he no he n egion
signi ican a 0.10 > P > 0.05.
Análisis de conglome ados mos ando las elaciones en e las localidades de mues eo a lo la go de la cos a del Pací ico de
Chile con inen al (escala egional). Las dis ancias es án basadas en los alo es semi-cuan i a i os de la abundancia de
cap élidos. Las líneas somb eadas indican ni eles de signi icancia: el g upo de es aciones de mues eo de la egión no e
ue signi ica i o a una p obabilidad 0,10 > P > 0,05.
THIEL ET AL.
305
ing communi y on buoys and opes e sus algal
and seag ass beds) sugges ing ha he cap ellid
species ound in he bay sys em o Coquimbo a e
ela i ely unselec i e wi h espec o hei subs a-
um, which appea s o be gene ally ue o li o al
cap ellid species. Fo example, Cap ella scau a
has been epo ed om b yozoans, sponges,
seaweeds and seag asses (Lim & Alexande 1986,
Takeuchi & Hino 1997, Se ejo 1998). Cap ella
equilib a has been collec ed om seaweeds, b yo-
zoans, sponges and ascidians (K app-Schickel
1993). Aoki & Kikuchi (1990) ound C. e ucosa
on e y di e en subs a a, and Cap ellina
longicollis has been collec ed om algae, b yozo-
ans and ascidians along he coas o New Zealand,
whe e i is he mos abundan species (McCain
1969). Maye (1890) epo ed ha C. e ucosa
and D. enenosa we e o en ound on a ious
subs a a belonging o he ouling communi y along
he coas o Coquimbo. In gene al, a he local
scale, he en i onmen al habi a condi ions (such
as hyd odynamics, sedimen a ion a e, u bidi y,
subs a um s abili y) seem o be mo e impo an in
de e mining he dis ibu ion o li o al cap ellids
han subs a um cha ac e is ics (Takeuchi e al.
1987, 1990, Gue a-Ga cía & Ga cía-Gómez 2001).
Recen ly, Gue a-Ga cía & Ga cía-Gómez (2001)
ound ha he species composi ion o he
Cap ellidea om one pa icula subs a um ( he
seaweed Cys osei a usneoides) along he coas o
Ceu a (No h A ica) changed conside ably ac-
co ding o en i onmen al ac o s such as local
hyd odynamics. This sugges s ha en i onmen al
condi ions a e usually mo e impo an in de e -
mining he dis ibu ion o li o al cap ellids han
subs a um cha ac e is ics. The impo ance o hy-
d odynamic condi ions o he dis ibu ion o li -
o al cap ellids is no su p ising since many spe-
cies eed di ec ly on o ganisms o ma e ials cap-
u ed om he wa e column (Takeuchi & Hi ano
1995).
Hyd odynamic condi ions may also ha e impli-
ca ions o cap ellid dispe sal wi hin bay sys-
ems. Mos bays along he con inen al coas o
Chile a e cha ac e ized by s ong ci cula ion and
e icien mixing o su ace wa e s (Soba zo 2002).
This con ibu es o e icien exchange and a ela-
i ely uni o m dis ibu ion pa e n o ben hic spe-
TABLE 2
Abundance o cap ellid species on de ached buoys o he Paci ic coas o Chile: p esen (1-10
indi iduals); common (11-100 indi iduals); abundan (101-1,000 indi iduals); supe abundan
(>1,000 indi iduals). All abundance es ima es ep esen minimum alues; (Cq) Coquimbo;
(Ca) Calde a. Fo de ails abou buoys see Appendix 1
Abundancia de las especies de cap élidos en boyas desp endidas y encon adas en e a la cos a Pací ico de Chile:
p esen e (1-10 indi iduos); común (11-100 indi iduos); abundan e (101-1.000 indi iduos); supe abundan e (>1.000
indi iduos). Todas las es imaciones de abundancia ep esen an alo es mínimos; (Cq) Coquimbo; (Ca) Calde a. Pa a
más de alles e Anexo 1
Coquimbo Calde a
Cq1 Cq2-b Cq2- Cq3 Cq4 Cq5 Cq6 Ca1 Ca2-b Ca2-
Sample ype Buoy Buoy Rope Buoy Buoy Buoy Buoy Buoy Buoy Rope
La i ude 29
o
49’23” 29
o
47’06” 29
o
58’38” 30
o
03’56” 30
o
13’00” 29
o
37’59” 27
o
01’23” 27
o
05’45”
Longi ude 71
o
26’39” 71
o
27’44” 71
o
24’59” 71
o
40’55” 71
o
42’35” 71
o
31’33” 70
o
56’55” 71
o
00’15”
Dis ance om sho e (km) 12.5 11.8 1.9 19.7 7.7 20 6.8 6.5
Cap ellina longicollis
Cap ella equilib a
Cap ella scau a
Cap ella e ucosa
Deu ella enenosa
Pa acap ella pusilla
Abundance pe sample:
P - p esen (1-10 indi iduals)
Co - common (11-100 indi iduals)
Ab - abundan (101-1,000 indi iduals)
Su - supe abundan (> 1,000 indi iduals)
DISTRIBUTION OF LITTORAL CAPRELLIDS FROM CHILE
312
APPENDIX 1
De ailed in o ma ion abou de ached buoys ound o he Paci ic coas o Chile: (Cq) Coquimbo; (Ca) Calde a; sampling olumes di e ed due
o logis ic easons (e.g., leng h o ope s ill a ached o buoys)
In o mación de allada de boyas desp endidas encon adas al en e de la cos a Pací ico de Chile: (Cq) Coquimbo; (Ca) Calde a; olumenes de mues eo di ie en po azones
logís icas (e.g., la go de la cue da aun ichado a las boyas)
Coquimbo Calde a
Cq1 Cq2-b Cq2- Cq3 Cq4 Cq5 Cq6 Ca1 Ca2-b Ca2-
Sample ype Buoy Buoy Rope (7m) Buoy Buoy Buoy Buoy Buoy Buoy Rope (12m)
Volume (1000 cm
3
) 11 21.5 11 11 11 11 11 21.5
Su ace a ea (100 cm
3
) 23.8 37.4 23.8 23.8 23.8 23.8 23.8 37.4
La i ude 29
o
49’23” 29
o
47’06” 29
o
58’38” 30
o
03’56” 30
o
13’00” 29
o
37’59” 27
o
01’23” 27
o
05’45” ..
Longi ude 71
o
26’39” 71
o
27’44” 71
o
24’59” 71
o
40’55” 71
o
42’35” 71
o
31’33” 70
o
56’55” 71
o
00’15” ..
Dis ance om sho e (km) 12.5 11.8 1.9 19.7 7.7 20 6.8 6.5
Sampling da e 11 June 2002 11 June 2002 12 June 2002 12 June 2002 12 June 2002 24 June 2002 9 Feb ua y 2002 9 Feb ua y 2002 ..
Sampling olume (l) 0.5 0.5 1.0 1.0 0.5 0.5 0.1 0.5 1.3
Buoy en i ely/pa ially sc aped En i e Pa ial En i e En i e En i e En i e En i e En i e Pa ial Pa ial
O he epibio a Ul a sp. B yozoa spp. Ul a spp., B yozoa spp., B yozoa spp. B yozoa spp. Ul a spp. Hyd ozoa spp. Hyd ozoa spp. Hyd ozoa spp.
B yozoa spp., B yozoa spp Ul a spp., Ul a spp.,
Ascidia spp. Bugula Bugula ne i ina,
ne i ina,Ascidia spp.
Ascidia spp. Pyu a chilensis
THIEL ET AL.