SCIENTIA MARINA 73(2)
June 2009, 399-407, Barcelona (Spain)
iSSn: 0214-8358
doi: 10.3989/scimar.2009.73n2399
Two new polychaete species living in the mantle cavity
of Calyptogena gallardoi (Bivalvia: Vesicomyidae) at a
methane seep site off central Chile (~36ºS)
Eduardo Quiroga 1 and JaviEr SEllanES 2,3
1 Centro
de investigación en Ecosistemas de la Patagonia (CiEP), Bilbao 449, Coyhaique, Chile.
E-mail: eduardo.quiroga@ciep.cl
Católica del norte, departamento de Biologia Marina, Facultad de Ciencias del Mar, larrondo 1281,
Coquimbo, Chile.
3 Centro de investigación oceanográica en el Pacíico Sur-oriental (CoPaS), universidad de Concepción, Casilla 160-C,
Concepción, Chile.
2 universidad
SuMMarY: Two new polychaete species belonging to nautiliniellidae and antonbruunidae were found in the mantle cavity of the vesicomyid bivalve Calyptogena gallardoi Sellanes and Krylova, 2005, at a methane seep site off central Chile.
Shinkai robusta n. sp. is characterized by having modiied parapodia with robust notopodia and nine simple hooks per parapodium on the middle setigers, and an anteriorly truncated sub-triangular prostomium, with a pair of small cirriform antennae.
The new species closely resembles Shinkai longipedata Miura and ohta, 1991, and Shinkai semilonga Miura and Hashimoto,
1996. Antonbruunia gerdesi n. sp. is characterized by having a trapezoidal prostomium, with ive sub-equal occipital antennae and a conspicuous pygidium with two short, well-developed digitiform anal cirri. These two new species constitute the
irst report of polychaetes living in symbiosis with chemosymbiotic bivalves in the south-eastern Paciic.
Keywords: methane seep, nautiliniellidae, antonbruunidae, symbiont polychaetes, Chile.
rESuMEn: Dos nuevas especies de poliquetos que habitan la cavidad del manto de
CALYPTOGENA GALLARDOI (Bivalvia: Vesicomyidae) en un afloramiento de metano frente a
Chile central (36°S). – dos nuevas especies de poliquetos simbiontes pertenecientes a las familias nautiliniellidae
y antonbruunidae se encontraron en la cavidad del manto del bivalvo vesicómyido Calyptogena gallardoi Sellanes y Krylova, 2005, proveniente de una zona de aloramiento de metano frente a Chile central. Shinkai robusta n. sp. se caracteriza por
presentar parapodios modiicados con notopodios bien desarrollados y robustos, nueve setas simples por parapodio en la
región media del cuerpo y un prostomio sub-triangular, truncado anteriormente, con un par de pequeñas antenas cirriformes.
la nueva especie es similar a Shinkai longipedata Miura y ohta, 1991, y Shinkai semilonga Miura y Hashimoto, 1996.
Antonbruuniia gerdesi n. sp. se caracteriza por presentar un prostomio trapezoidal, con cinco antenas occipitales sub-iguales, incluyendo una antena central bien desarrollada, insertada en la supericie dorsal posterior del prostomio, y un pigidio
conspicuo con dos cirros anales cortos digitiformes. Estas dos nuevas especies constituyen la primera cita de poliquetos
simbiontes de bivalvos quimiosimbióticos en el Pacíico sur-oriental.
Palabras clave: aloramiento de metano, nautiliniellidae, antonbruunidae, poliquetos simbiontes, Chile.
inTroduCTion
Benthic communities thriving in reducing habitats such as methane seeps and hydrothermal vents
are constituted by rich invertebrate assemblages
that host chemoautotrophic symbionts (e.g. Sibuet
and olu, 1998; Sibuet and olu-le roy 2002; Sahling et al., 2002; van dover et al., 2003), as well as
many heterotrophic species that also beneit from the
chemosynthesis and methane-derived carbon (Bis-
400 • E. Quiroga and J. SEllanES
coito et al., 2002; van dover et al., 2003; levin,
2005; Cordes et al., 2007). Communities of benthic
animals associated with methane seeps (also known
as cold seeps) have been found in several locations
on active and passive continental margins of the
Paciic and atlantic ocean (olu et al., 1996; olule roy et al., 2004; ravara et al., 2007; Cordes et
al., 2007). There have been some recent studies on
the occurrence of cold seepage and gas hydrates on
the central Chile margin (Morales, 2003; Sellanes
et al., 2004, 2008). Clams dominate the chemosymbiotic fauna at a cold seep site located off central
Chile (36ºS) called the Concepción Methane Seep
area (CMSa) (Sellanes et al., 2004; Sellanes and
Krylova, 2005). The chemosymbiotic assemblages
at the CMSa include vesicomyid, solemyid, lucinid
and thyasirid bivalves (Sellanes and Krylova 2005;
Holmes et al., 2005; oliver and Sellanes, 2005), as
well as a siboglinid tubeworm of the genus Lamellibrachia (Sellanes et al., 2008).
vesicomyid bivalves are typical organisms of marine chemosynthesis-based communities (e.g. Sibuet
and olu-le roy, 2002; van dover et al. 2003; Sellanes et al., 2008). in the CMSa, the most frequently
collected species is Calyptogena gallardoi, and two
polychaete species belonging to the families nautiliniellidae (Miura and laubier, 1989) and antonbruunidae Fauchald, 1977 were found within its mantle cavity. nautiliniellids are a small group of polychaetes
and all the species reported so far live as symbionts
within deep-sea bivalve molluscs from hydrothermal
vents and cold seeps (e.g. Miura and laubier, 1989,
1990; Blake, 1993; Miura and Hashimoto, 1996;
dreyer et al., 2004; ravara et al., 2007). This family
has eleven genera, which are characterized by having
a muscularized foregut, paired antennae, simple neuropodial hooks, and a rounded pygidium without anal
cirri (Blake, 1993; dreyer et al., 2004). However, the
taxonomic status and ecology of this family are still
not fully understood (Blake, 1993, 1997). The family
antonbruunidae was recognized by Fauchald (1977),
and is closely related to the family nautiliniellidae
(Martin and Britayev, 1998). Antonbruunia viridis
Hartmann and Boss, 1965, the only species described
so far, is associated with the shallow water (62 to 82
m depth) bivalve mollusc Lucina fosteri Hartmann
and Boss, 1965.
in fact, since both families are morphologically
very close, it has been suggested that their taxonomic position should be redeined (see Martin and
Britayev, 1998). The two new species described
in this paper constitute the irst report of symbiont
polychaetes of chemosymbiotic bivalves in the
south–eastern Paciic.
MaTErialS and METHodS
Study site
The site is located 72 km nW off Concepción
Bay, Chile, at the slope zone (750 to 900 m water
depth) and near a mound separated by a shallow
depression from another mound (Fig. 1). The pres-
Fig. 1. – Study site, located on the slope zone off central Chile (36ºS). The triangle indicates the area in which trawls successfully collected
evidence of active methane seepage (carbonate blocks, live chemosymbiotic clams and shell fragments). The star indicates the locations where
shallow sub-surface gas hydrates have been observed.
SCi. Mar., 73(3), June 2009, 399-407. iSSn 0214-8358 doi: 10.3989/scimar.2009.73n2399
SYMBionT PolYCHaETES oF CALYPTOGENA • 401
Fig. 2. – Shinkai robusta n. sp. (agT10/vg-07): a, anterior region, ventral view; B, prostomiun, ventral view; C, anterior region, lateral view;
d, parapodia of the middle region; E, hooks of anterior parapodia (arrow showing a stout hook); F, pygidium, dorsal view.
ence of mounds and pockmarks has been previously documented for areas rich in subsurface gas
hydrate deposits (dando et al., 1991; Sassen et al.,
2001, 2003). The sediment surface is characterized
by the presence of abundant carbonate-cemented
mud blocks (Sellanes et al., 2004, 2008), while
high concentrations of methane and sulide, as well
as gas hydrates, have been documented in sediment
cores (Cofin et al., 2006). Chemosymbiotic bivalves include vesicomyids, the solemyid Acharax
sp., the lucinid Lucinoma anemiophila, and the
thyasirids Thyasira methanophila and Conchocele
sp. (Holmes et al., 2005; oliver and Sellanes, 2005;
Sellanes et al., 2008).
Collection of samples
Polychaete specimens were collected during oceanographic cruises conducted onboard rv “vidal
gormáz” of the Chilean navy during october 2004
(vg-04 cruise), September 2006 (Seepox cruise),
and September 2007 (vg-07 cruise). The biological
material was obtained with an agassiz trawl (mouth
opening 1.5 x 0.4 m, mesh size 10 x 10 mm at the
cod-end) in 20 minute hauls.
Specimens of Calyptogena gallardoi were dissected onboard immediately after collection and
polychaetes were extracted and ixed in 10% formalin and later preserved in 70% ethanol. details of
SCi. Mar., 73(3), June 2009, 399-407. iSSn 0214-8358 doi: 10.3989/scimar.2009.73n2399
402 • E. Quiroga and J. SEllanES
Fig. 3. – Shinkai robusta n. sp. (agT10/vg-07): a, anterior region, ventral view; B, anterior region, lateral view; C, neuropodial hooks from
parapodium 6; d, pygidium, dorsal view; E, parapodium 6 of the holotype; F, middle parapodium.
the setae were examined under immersion oil with
a compound microscope. The terminology used follows Miura and laubier (1989, 1990), Miura and
ohta (1991), Miura and Hashimoto (1993, 1996)
and Blake (1990, 1993). Scanning electronic microscope (SEM) photographs were obtained (JEol
JSM-T300 microscope) of critical-point dried, gold
coated specimens (JEol JFC-1100 ine coat ion
sputter; rouse and Pleijel, 2001).
Type specimens are deposited in the “Museo nacional de Historia natural”, Santiago, Chile (MnHnCl) and the reference collection of the Centre for
advance Studies in Patagonian Ecosystems (CiEP),
Coyhaique, Chile.
SYSTEMaTiC aCCounT
Family Nautiliniellidae (Miura and laubier, 1989)
genus Shinkai Miura and laubier, 1990
Shinkai robusta n. sp.
(Figs. 2 and 3)
Type material. Holotype (MnHnCl-an2043), complete
specimen, collected from the mantle cavity of Calyptogena
gallardoi. Type locality: off Concepción, Chile (36°21’64”S,
73°43’57”W), 865-926 m deep. october, 11, 2004 (agT7/ vg04). Paratype (1 specimen, used for SEM photographs), complete specimen, collected from the mantle cavity of Calyptogena
gallardoi. Type locality; off Concepción, Chile (36°22’01”S,
73°43’10”W), 764-843 m deep. october 02, 2007 (agT10/vg07).
Description. (Holotype); complete specimen
with 78 segments, measuring 13.86 mm long, 2.13
mm wide, including parapodia. Paratype; complete
specimen with 82 setigers, measuring 19.20 mm
long, 2.32 mm wide, including parapodia. Body
elongated, vermiform, lattened ventrally, strongly
arched dorsally and with longitudinal ventral groove.
Colourless in alcohol. Prostomium short, sub-triangular with pair of small cirriform antennae, without
eyes or other appendages. Mouth opening ventrally
between prostomium and irst setiger (Fig. 2a-C,
3a-B). Foregut with well-developed muscular region
without jaws and paragnaths. First segment partially
fused with prostomium. achaetous peristomial ring
absent (Fig. 2C). Parapodia subbiramous with welldeveloped dorsal cirri and reduced ventral cirri (Fig.
2d-E and 3a-B, E-F). neuropodia well developed,
supported by single, stout neuroacicula; ventral cirri
shorter than dorsal cirri (Fig. 3E, F). anterior neuropodia with simple hooks (Fig. 3C), six on parapodia
SCi. Mar., 73(3), June 2009, 399-407. iSSn 0214-8358 doi: 10.3989/scimar.2009.73n2399
Mozambique Channel Hartmann and Boss, 1965
Central Chile, CMSa Present study
Lucina fosteri Hartman and Boss, 1965 (lucinidae)
Calyptogena gallardoi Sellanes and Krylova, 2005 (vesicomyidae)
Family antonbruunidae
Antonbruunia viridis Hartman and Boss, 1965
Antonbruunia gerdesi n. sp
68-82
795-843
Blake, 1993
Miura and Hashimoto, 1996
Blake, 1993
Blake, 1993
Miura and Hashimoto, 1993
Miura and Hashimoto, 1993
Miura and laubier, 1990
Miura and Hashimoto, 1996
Miura and laubier, 1989
Blake, 1990
Miura and Hashimoto, 1996
dreyer et al., 2004
Miura and otha, 1991
Miura and laubier, 1990
Miura and Hashimoto, 1996
Present study
Florida Escarpment
okinawa Trough
Florida Escarpment
Santa Maria Basin
okinawa Trough
okinawa Trough
Sagami Bay
Kagoshima Bay
Japan Trench
laurentian Fan
Sagami Bay
Blake ridge diapir
okinawa Trough
Sagami Bay
okinawa Trough
Central Chile, CMSa
unknown
Bathymodiolus aduloides Hashimoto and okutani, 1994 (Mytilidae)
Mytilidae
unknown
Bathymodiolus sp. (Mytilidae)
Bathymodiolus aduloides Hashimoto and okutani, 1994 (Mytilidae)
Solemya sp. (Solemyidae)
Solemya sp. (Solemyidae)
Calyptogena phaseoliformis Mètivier et al., 1986 (vesicomyidae)
Thyasira insignis (verrill and Bush, 1898)(Thyasiridae)
Conchocele disjuncta gabb, 1866 (Thyasiridae)
Calyptogena sp. (vesicomyidae)
Calyptogena sp. (vesicomyidae)
Calyptogena soyoae okutani, 1957 (vesicomyidae)
Calyptogena solidissima okutani et al. 1992 (vesicomyidae)
Calyptogena gallardoi Sellanes and Krylova, 2005 (vesicomyidae)
Family nautiliniellidae
Flascarpia alvinae Blake, 1993
Iheyomytilidicola tridentatus Miura and Hashimoto, 1996
Laubierus mucronatus Blake, 1993
Miura spinosa Blake, 1993
Mytilidiphila enseiensis Miura and Hashimoto, 1993
Mytilidiphila okinawaensis Miura and Hashimoto, 1993
Natsushima bifurcata Miura and laubier, 1990
Natsushima graciliceps Miura and Hashimoto, 1996
Nautiliniella calyptogenicola (Miura and laubier, 1989)
Petrecca thyasira Blake, 1990
Thyasiridicola branchiatus Miura and Hashimoto, 1996
Vesicomyicola trifurcatus dreyer et al., 2004
Shinkai longipedata Miura and otha, 1991
Shinkai sagamensis Miura and laubier, 1990
Shinkai semilonga Miura and Hashimoto, 1996
Shinkai robusta n. sp.
3303
1395
3243
565
625
701
1114
98
5650
3700
1160
2155
1400
1170
625
764-926
location
depth (m)
Host bivalve
Habitat. all previously known nautiliniellid species are symbionts associated with bivalve molluscs
from cold seeps or hydrothermal vents (Table 1).
The genus Shinkai is close to Nautiliniella, but differs in having up to eight hooks per neuropodium
instead of one, and a single pair of antennae instead
of two. in contrast, Natsushima differs from the
other two genera as it has two types of neuropodial
spines instead of only one (Blake, 1993). The genus
Shinkai comprises, at present, three described species: Shinkai longipedata Miura and otha, 1991,
Shinkai semilonga Miura and Hashimoto, 1996 and
Shinkai sagamensis Miura and laubier, 1990, all
from Japan. The occurrence rate in bivalves of S.
Species
Remarks. The new species closely resembles
the other congeneric species. in general, the parapodia and the head region of Shinkai robusta n. sp
are similar to those of the Shinkai semilonga and
Shinkai longipedata specimens; however, the main
diagnostic characters are the shape of the neuropodial hooks, their number on each parapodium and
the sub-triangular prostomium with a pair of small
cirriform antennae. in fact, S. robusta n. sp. differs
from all other species of the genus as it has anterior
neuropodia with six hooks on parapodia 1-10, and
one of the hooks is stouter than the others. in S. semilonga, the head and irst eight setigers are depressed
antero-posteriorly. The neuropodial hooks are simple
and slightly curved at the distal end. The number of
hooks per parapodium is about 15 on parapodia 1-3,
and more than 25 on parapodia 4-6, then decreases
to about ten on parapodium 10, and then ive to eight
on parapodia 50 to 200 (Miura and Hashimoto, 1996;
Fig. 4a-E). in S. longipedata the middle and posterior parapodia are armed by a single neuropodial hook
with a strongly curved distal fang (Fig. 4F-i; Miura
and Hashimoto, 1996; dreyer et al., 2004). Shinkai
sagamensis differs from the above three species as
it has very short notopodia and a different number
of hooks on each parapodium (Fig. 4J-l). in fact,
several hooks projected from each neuropodium on
the anterior parapodia, e.g. 3-4 on parapodium 1, 5-8
on parapodia 2-6, 1-3 on parapodia 7-20 and 1 on the
posterior parapodia (Miura and laubier, 1990).
Table 1. – Families nautiliniellidae and antonbruunidae: list of species, host bivalve species, collection depth, location and author references.
1-10, one of the hooks is stouter than the others (see
Fig. 2C, E and 3C, E, F). Posterior neuropodia with
up to nine simple hooks per neuropodium, all of
them similar (Fig. 3F). Pygidium rounded, without
anal cirri (Fig. 2F, 3d).
references
SYMBionT PolYCHaETES oF CALYPTOGENA • 403
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404 • E. Quiroga and J. SEllanES
Fig. 4. – Shinkai semilonga (after Miura and Hashimoto, 1996): a, anterior region, dorsal view; B, anterior region, lateral view; C, pygidium,
dorsal view; d, parapodium 100; E, neuropodial hooks. Shinkai longipedata (after Miura and otha, 1991): F, anterior region, dorsal view; g,
pygidium, dorsal view; H, neuropodial hook; i, parapodium 100. Shinkai sagamensis (after Miura and laubier, 1990): J, anterior region, dorsal
view; K, pygidium, dorsal view; l, neuropodial hook.
robusta n. sp. is very low, and only two of the 35
specimens of C. gallardoi analyzed hosted a single
individual of this polychaete species.
Distribution. only known from the type locality
off Concepción, at a depth of 764 to 926 m. Shinkai
robusta n. sp. is the irst nautiliniellid species reported at a cold seep site in the south-eastern Paciic
area. To date, its host bivalve C. gallardoi is also
only known from the type locality.
Etymology. The species’ name is derived
from their characteristically elongated and robust
notopodia.
Family Antonbruunidae Fauchald, 1977
genus Antonbruunia Hartman and Boss, 1965
Antonbruunia gerdesi n. sp.
(Figs. 5 and 6)
Type material. Holotype (MnHnCl-an2044), complete specimen, collected from the mantle cavity of Calyptogena gallardoi.
Type locality; off Concepción, Chile (36°22’19”S, 73°43’36”W),
843-846 m deep. 04 September, 2004 (agT6-9/Seepox). Paratype
(1 specimen, used for SEM photographs), complete specimen, collected from the mantle cavity of Calyptogena gallardoi. Type locality; off Concepción, Chile (36°21’93”S, 73°42’84”W), 795-843 m
deep. 02 September, 2006 (vg-06/Seepox).
Description. (Holotype); complete specimen
with 33 segments, measuring 5.86 mm long, 1.10
mm wide, including parapodia. Paratype; complete
specimen with 53 segments, measuring 16.40 mm
long, 2.04 mm wide, including parapodia. Body vermiform, lattened ventrally, slightly arched dorsally,
without longitudinal ventral groove (Fig. 5a-C, and
Fig 6a). Colour green in life, white after preservation.
Prostomium short and trapezoidal with ive sub-equal
cirriform antennae, including an unpaired median
antenna, without eyes or other appendages. Frontal
antennae shorter, inserted ventrally on prostomium
(Fig. 5a, B and 6B). Median antenna inserted posteriorly on the dorsal surface of the prostomium. Pharynx
without jaws, paragnaths. Peristomial achaetous ring
with two pairs of long, well-developed cirri (Fig. 5a,
B and 6B). Parapodia increasing gradually to the postanterior and mid-body region (Fig. 5a). all parapo-
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SYMBionT PolYCHaETES oF CALYPTOGENA • 405
Fig. 5. – Antonbruunia gerdesi n. sp. (agT6-9/Seepox): a, anterior region, ventral view; B, anterior region, dorsal view; C, parapodia of the
middle region lateral view; d, parapodia of the middle region, ventral view, E, hooks of the middle region; F, hooks; g, pygidium, ventral
view.
dia subbiramous (Fig 5C-E and 6C). notopodia with
dorsal cirrus and a slender embedded notoaciculum,
straight and pointed. all setae simple with long
straight shaft, biid, with two straight teeth of similar
length (Fig. 5E, F and 6B, d). Pygidium conspicuous,
with two well-developed digitiform cylindrical anal
cirri (Fig. 5g, 6C).
Remarks. Antonbruunia gerdesi n. sp. differs
from A. viridis in the shape of its notopodia (elongated and wide), and the presence of ive sub-equal
occipital antennae on the prostomium (Fig. 6a-d,
6F-H). The hooks of A. gerdesi n. sp are simple, with
a long, straight, distally biid shaft (Fig. 6E). in contrast, those of A. viridis are curved distally (Miura
and laubier, 1990; Fig. 6H). The only known species that represents the family antonbruunidae so
far, Antonbruunia viridis inhabits the mantle cavity
of its host bivalve Lucina fosteri (Table 1), which
occurs in hypoxic sediments of black-brown oozy
mud and detritus, off the coast of Mozambique, indian ocean (Hartman and Boss, 1965).
Habitat. The host bivalve of A. gerdesi n. sp.
(C. gallardoi) is only known from the type locality in the south-eastern Paciic (Sellanes and
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406 • E. Quiroga and J. SEllanES
Fig. 6. – Antonbruunia gerdesi n. sp. (agT6-9/Seepox): a, photography of whole body, lateral view; B, anterior region, dorsal view; C,
middle parapodium; d, pygidium, ventral view; E, hook. Antonbruunia viridis Hartman and Boss, 1965 (after Miura and laubier, 1990): F,
anterior region, lateral view; g, pygidium, lateral view; H, posterior parapodium; i, hook (after Hartman and Boss, 1965).
Krylova, 2005), where it inhabits sticky dark grey
sediments smelling of sulphide. Carbonate crusts
also occur in the area, and gas hydrates were retrieved from subsurface sediments nearby. like
the other species described in the present study,
the occurrence rate of A. gerdesi n. sp. is low:
four of the 35 specimens of C. gallardoi analyzed
hosted an individual of this polychaete. although
in general each bivalve hosted a single polychaete,
three specimens were found in one host bivalve
that measured 9 mm long. it is also worth noting
that no co-occurrence of A. gerdesi n. sp. and S.
robusta n. sp. has been observed within a single
specimen of C. gallardoi, but to our knowledge
this is the irst time that two different polychaete
species have been found associated with the same
vesicomyid host species.
Distribution. only known from the type locality off
Concepción (Chile), 795-846 m depth. Antonbruunia
gerdesi n. sp. is the second known species of the family, the irst antonbruunid found in the Paciic ocean,
and the irst report of the family at a cold seep site.
Etymology. The species is named in honour of
dr. dieter gerdes (alfred Wegener institute for Polar and Marine research, germany) who has studied
marine benthic communities of the South american
and antarctic margins extensively.
aCKnoWlEdgEMEnTS
our thanks go to the oficers and crew of r/v vidal
gormáz for their skilful operations at sea. We thank
Juan diaz naveas (PuCv, Chile) for providing onboard
facilities during the vg-04 and vg-07 cruises. Taxonomic literature was provided by dr. dieter gerdes
(aWi, germany) and dr. guillermo San Martin (uaM,
Spain). We are also very grateful to Maria Soledad
romero, universidad Católica del norte (Coquimbo)
for help in the preparation of the specimens and SEM
work. We thank Paula ortiz Saini for her patience and
dedication in preparing the drawings. The manuscript
beneited from reviews by Jennifer dreyer (virginia
institute of Marine Science, uSa), guillermo guzmán
(universidad arturo Prat, Chile), dr. américo Mon-
SCi. Mar., 73(3), June 2009, 399-407. iSSn 0214-8358 doi: 10.3989/scimar.2009.73n2399
SYMBionT PolYCHaETES oF CALYPTOGENA • 407
tiel (universidad de Magallanes, Chile) and insightful
comments provided by dr. Tomoyuki Miura and an
anonymous reviewer. This research was supported by
the FondECYT #1061217 project. The FondECYT
#1061214 and FondaP-CoPaS #15010007 projects
contributed extra funding for shiptime. We thank the
CiEP center (gobierno regional de aysen, Chile, BiP,
no. 3004258-0) for their support during the writing
phase of this work.
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received March 5, 2008. accepted october 17, 2008.
Published online March 12, 2009.
SCi. Mar., 73(3), June 2009, 399-407. iSSn 0214-8358 doi: 10.3989/scimar.2009.73n2399