Electronic Supplementary Information to the Manuscript
Bone-eating worms from the Antarctic: the contrasting fate of whale and wood remains on the
Southern Ocean seafloor
Adrian G. Glover1, Helena Wiklund1, Sergio Taboada2, Conxita Avila2,3, Javier Cristobo4, Craig R. Smith5,
Kirsty M. Kemp1,6, Alan Jamieson7, Thomas G. Dahlgren8,9*
1
Life Sciences Department, The Natural History Museum, Cromwell Road, London, UK, SW7 5BD
2
Depto. de Biología Animal, Facultad de Biología, Universidad de Barcelona, Avda. Diagonal 643, 08028
Barcelona, Spain
3
Biodiversity Research Institute, Campus Sud, Av. Diagonal 643, 08028 Barcelona, Spain
4
Centro Oceanográfico de Gijón, Instituto Español de Oceanografía, Avda. Príncipe de Asturias 70 bis,
33212 Gijón, Spain
Department of Oceanography, University of Hawai’i at Manoa, 1000 Pope Road, Marine Science Building,
Honolulu, HI96822 USA
5
6
Institute of Zoology, Zoological Society of London, Regent’s Park, London, UK, NV1 4RY
7
Oceanlab, University of Aberdeen, Institute of Biological and Environmental Sciences,
Main Street, Newburgh, Aberdeenshire, UK, AB41 6AA
8
University of Gothenburg, Department of Biological and Environmental Sciences, P.O. Box 463, 405 30
Gothenburg, Sweden
9
Present address: Uni Research, P.O. Box 7810, 5020 Bergen, Norway
*correspondence: thda@mac.com
Detailed Materials and Methods
Study area and experimental setup
The study area and sites were chosen based on two criteria: how representative they are of the typical
Antarctic shelf habitat, and the logistical fit to pre-existing oceanographic cruise programs to minimise cost.
Free-vehicle landers from the Antarctic Chemosynthetic Ecosystems (ACES) project were used at two
locations (ACES 1 and 2, near to Smith Island), and a simple mooring on an acoustic release at the site in
Whalers Bay, Deception Island (Main Text, figure 1a,b). The site near Smith Island lies on a relatively flat
area of soft-sediment shelf environment at 550-650m depth close to the 'FOODBANCS2 Project' site 'AA',
approximately 30 nautical miles south-west of Deception Island (Smith et al. 2011). The site in Whalers Bay,
Deception Island, lies at the much shallower depth of 21m, close to the early 20th century whaling station.
Whalers Bay is a highly unusual benthic environment, disturbed by the presence of geothermal activity
(causing an influx of warm water into the harbour) (Somoza et al. 2004) and with a large number of whale
bones and debris discarded by the whalers during the period 1906-1931 (Tønnessen and Johnsen 1982).
SCUBA and ROV surveys in 2005, 2009, 2010 and 2011 have since confirmed the presence of a very large
number of whale bones on the seafloor at depths of 0-60m in Whalers Bay [T.G.D, J.C, C.A., S.T. pers obs.,
Jon Copley, pers. comm.]
This is the first time that a free-vehicle lander has been successfully modified for the study of deep-sea whale
and wood-falls. The ACES experiments utilised a modified compact lander supplied by Oceanlab, University
of Aberdeen. Each lander consisted of an aluminum frame surrounded by fixed-buoyancy foam blocks, a
lifting ring, flag, separate surface float and radar reflector to aid surface recovery, ballast and a 6000m-rated
IXSEA Acoustic release unit (Main Text, figure 1c). Each lander was equipped with experimental settlement
substrates consisting of minke whale (Balaenoptera acutorostrata Lacépède, 1804), oak wood (Quercus
robur L.) and pine wood (Pinus sylvestris L.). The whale bones for ACES 1 and 2 were collected from a
dead, stranded carcass in Fjällbacka, Sweden on 4th April 2007, de-fleshed, sterilized in freezers and
transported frozen aboard the MS Oden to Antarctica. The oak and pine wood was collected in eastern
Sweden. Minke whale bones were chosen as they are abundant in the Antarctic, in the form of the species
Balaenoptera bonaerensis Burmeister, 1867 (morphologically almost identical to B. acutorostrata). Oak and
pine planks from Scandinavia were used as these were the building materials for the wooden ship Endurance
(now a shipwreck in the Antarctic), as well as other early 20th century shipwrecks, e.g. Otto Nordenskjöld's
Antarctic.
For the ACES 1 experiment, the substrates consisted of 1 skull, 2 vertebra, 3 bone pieces, 8 pine planks and
1 oak plank, with a total weight of 93.5 kg. For the ACES 2 experiment, the substrates consisted of 1
scapula, 3 vertebra, 1 rib, 4 bone pieces, 8 pine planks and 1 oak plank with a total weight of 24.2 kg. The
Whalers Bay mooring consisted of a single caudal vertebra (approximately 20cm in length, width and height)
of a juvenile B. acutorostrata collected from a dead stranding in Asturias, Spain and treated in a similar
manner to the ACES bones. The mooring itself was simply the bone attached to a piece of ballast, with a line
going to an IXSEA 500 release, flotation and a larger piece of ballast. No wood was attached to the Whalers
Bay mooring.
The ACES 1 and 2 experiments were deployed by the Swedish Icebreaker MS Oden and the Whalers Bay
mooring by a vessel from the Spanish Research Station ‘Gabriel de Castilla’ in December 2007 and January
2009 respectively (Main Text, table 1). The ACES experiments were recovered by acoustic command (and
visual search) in February 2009 by the US vessel RV Laurence M Gould and the Whalers Bay mooring was
recovered in January 2010 by the Spanish scientists. Thus each mooring had between 12-14 months on the
seafloor.
For both the ACES lander experiments, wood and whale-bone were combined onto the same mooring in
order to maximise sample return per effort. We assessed the potential caveat that having two differing
substrates on the same landers may influence the larval settlement, for example the presence of whale bones
could inhibit larval settlement by bivalves. However, we currently discount this based on data from
experimental deployments at shelf depths in the North Sea, Sweden and bathyal depths in the central Indian
ocean, where bones and wood were deployed together with no observed inhibition of bivalve larval
settlement on the wood (T Dahlgren, K Kemp, A Glover, unpublished data).
Sampling
On recovery, the bones and wood from the ACES experiments were detached from the lander frame and
stored in onboard aquarium facilities at ambient temperature of 1-2°C prior to processing. Photographs of the
substrate surface and fauna were taken underwater using an Olympus underwater camera. Specimens were
picked and (in the case of Osedax) dissected from the whale bone under stereo microscopes and imaged
using a Nikon 4500 camera with microscope adaptor (for ACES sites) and Invenio 5S CMOS with Zeiss
Stemi-2000 (for Whalers Bay site) before preservation. Material that had washed off the substrates and
collected in the bottom of the tanks was sieved at 300µm and examined under microscope. The majority of
the fauna was later preserved in 95% ethanol in individual numbered vials, with some specimens selected for
preservation in 10% formalin or 2% glutaraldehyde in sodium cacodylate buffer for more detailed
morphological study.
For the material from Whalers Bay, a similar protocol was used although no formalin-fixation was carried
out. The bone was brought to the laboratory at the Spanish Antarctic Base 'Gabriel de Castilla' and placed in
an aquarium maintained at room temperature (0-5°C) with no oxygenation of the water.
Morphological analysis
In the laboratory, all specimens were databased and identified to putative species level. Specimens were
examined and imaged using light and electron microscopy (SEM). Light microscopy was undertaken using
Wild M5 field microscopes equipped with a Nikon Coolpix 4500 camera, a Leica DM5000 compound
microscope with DFC 480 camera and a Zeiss V.20 stereomicroscope with AxioCam camera. For SEM,
specimens were critical-point-dried, gold-coated and imaged using a Philips XL-30 FEG scanning electron
microscope. The individual males of Osedax antarcticus sp. nov. (approximately 500µm in length) were
recovered separately from the tubes of the females, and critical-point-dried on their own. They were then
collected from the drying chamber baskets using an eyelash attached to a stick, and carefully placed
transversely across a piece of razor blade coated with a thin veneer of epoxy resin, orientated vertically
above the stub, allowing the majority of the specimen, including the chaetae, to be imaged. The internal
bacterial symbionts of the root tissue were imaged through a transverse section of root tissue, with exposed
tissue regions gold-coated and SEM imaged as above.
38 formalin-preserved specimens of female Osedax antarcticus sp. nov. from the Smith Island site were
examined, imaged and measured using the Zeiss V.20 and AxioCam camera. Measurements were taken of
the maximum depth and width of the roots, trunk, palps and oviduct. The number of dwarf males was
counted on each specimen. A futher 73 specimens of Osedax sp. indet. (hypothesised to be Osedax
antarcticus sp. nov.) were examined from a photo taken of live animals emergent from bone samples, the
maximum length of the trunk and palps was measured and analysed using Microsoft Excel.
Molecular sequencing and analysis
The molecular phylogenetic analyses were made with datasets from the sequences 18S, 16S and cytochrome
c oxidase subunit I (COI). In total, 46 terminal taxa were included in the analyses, 26 from Osedax, 17 from
other genera within Siboglinidae, and three outgroup taxa of which a spionid, Malacoceros fuliginosus
(Claparède, 1870), was used as root. Outgroup choice was based on analyses of Rousset et al. (2004) and
Struck et al. (2007). Accession numbers for the taxa used can be found in Table 1. In the Sabellidae
outgroup, 18S and 16S from Sabella pavonina Savigny, 1822 was used together with COI from Sabella
spallanzanii (Gmelin, 1791). In the Oweniidae outgroup, 18S and COI from Owenia fusiformis Delle Chiaje,
1844 was used together with 16S from Myriochele sp. Extraction of DNA was done with DNAeasy Tissue
Kit (Qiagen) following the protocol supplied by the manufacturer. About 1900 bp of 18S, 500 bp of 16S and
600 bp of COI were amplified. PCR mixtures contained ddH2O, 1 µl of each primer (10µM), 2 µl template
DNA and puReTaq Ready-To-Go PCR Beads (GE Healthcare) in a mixture of total 25 µl. The temperature
profile was as follows: 96ºC/240s -(94ºC/30s-48ºC/30s-72ºC/60s)*45cycles-72ºC/480s. PCR products were
purified with the E.Z.N.A. Cycle-Pure Kit (Omega Bio-tek). Sequencing was performed by the Macrogen
Sequencing System in Korea, on an ABI 3730XL DNA Analyser (Applied Biosystems), using primers listed
in Table 3.
Overlapping sequence fragments were merged into consensus sequences using Geneious (Kearse et al. 2012)
and aligned using MUSCLE (Edgar 2004) for COI, and MAFFT (Katoh et al. 2002) for 18S and 16S, both
alignment programs provided as plug-ins in Geneious and used with default settings. Bayesian phylogenetic
analyses (BA) were conducted with MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003). Analyses were run
three times for each dataset with four chains for 10,000,000 generations. 2,500,000 generations were
discarded as burn-in. The treefiles were tested with AWTY (Are We There Yet) (Nylander et al 2008) to see
that the analyses had reached a stationary phase. The evolutionary models used for the molecular data in BA
were obtained by running the datasets in MrModelTest (Nylander 2004), and for 18S the optional model was
GTR+I+G, while GTR+G was suggested for 16S. For COI the data was partitioned into codon positions, and
position 1 and 2 followed GTR+I+G, while HKY+G was used for position 3. A separate COI dataset was
made for haplotype distribution analysis using TCS 1.21 (Clement et al. 2000), with twelve sequenced
specimens of Osedax antarcticus sp. nov.
Results
ESM Table 1. Taxa, collection sites and NCBI GenBank accession numbers.
Terminal taxa
Alaysia sp. Southward, 1991
Escarpia spicata Jones, 1985
Galathealinum brachiosum Ivanov, 1961
Lamellibrachia barhami Webb, 1969
Lamellibrachia columna Southward, 1991
Lamellibrachia satsuma Miura, 1997
Malacoceros fuliginosus (Claparède, 1870)
Myriochele sp. Malmgren, 1867
Oasisia alvinae Jones, 1985
Oligobrachia haakonmosbiensis Smirnov, 2000
Osedax antarcticus sp. n.
Osedax deceptionensis sp. n.
Osedax frankpressi Rouse et al., 2004
Osedax japonicus Fujikura et al., 2006
Osedax mucofloris Glover et al., 2005
Osedax roseus Rouse et al., 2008
Osedax rubiplumus Rouse et al., 2004
Osedax sp. 'green palp'
Osedax sp. 'nude palp A'
Osedax sp. 'nude palp B'
Osedax sp. 'nude palp C'
Osedax sp. 'nude palp D'
Osedax sp. 'nude palp E'
Osedax sp. 'nude palp F'
Osedax sp. 'orange collar'
Osedax sp. 'pinnules'
Osedax sp. 'sagami 3'
Osedax sp. 'sagami 4'
Osedax sp. 'sagami 5'
Osedax sp. 'sagami 6'
Osedax sp. 'sagami 7'
Osedax sp. 'sagami 8'
Osedax sp. 'spiral'
Osedax sp. 'white collar'
Osedax sp. 'yellow collar'
Osedax sp. 'yellow patch'
Owenia fusiformis Delle Chiaje, 1844
Paraescarpia sp. Southward et al., 2002
Polybrachia sp. Ivanov, 1952
Ridgeia piscesae Jones, 1985
Riftia pachyptila Jones, 1981
18S
FM995545
AF168741
AF168738
AF168742
FJ347679
FM995543
AY525632
--AF168743
AM883186
KF444420
KF444421
FJ347682
FM995535
AY941263
FM995536
FM995538
FJ347694
FJ347687
FJ347686
FJ347688
FJ347691
FJ347692
FJ347695
FJ347690
--FM995537
FM995541
FM995539
FM995540
FM995542
FM995534
FJ347693
FJ347684
FJ347689
FJ347685
AB106256
FM995546
AF168739
AF168744
AF168745
16S
--AF315041
AF315040
AF315047
FJ347646
--EF431962
AY340468
AF315052
--KF444418
KF444419
AY577876
--FJ347657
AY577878
FJ347655
FJ347653
FJ347652
FJ347650
FJ347649
FJ347648
FJ347651
FJ347661
--------------FJ347647
FJ347659
FJ347660
FJ347654
--AF315053
AF315037
AF315054
AF315049
COI
--FJ667537
U74066
AY129146
DQ996645
D38030
EF432016
--AY646020
FM178481
KF444422
KF444428
DQ996621
FM998111
AY827568
EU032469
EU223302
FJ347640
EU223356
EU236218
EU267675
FJ347630
FJ347634
FJ347643
EU223340
FJ431197
FM998078
FM998082
FM998083
FM998091
FM998108
FM998110
FJ347637
FJ347611
EU223323
FJ347618
AY428839
D50595
FJ480393
AF022233
FJ667529
Sabella pavonina Savigny, 1822
Sabella spallanzanii (Gmelin, 1791)
Sclerolinum brattstromi Webb, 1964
Sclerolinum contortum Smirnov, 2000
Siboglinum ekmani Jägersten, 1956
Spirobrachia sp. Ivanov, 1952
Tevnia jerichonana Jones, 1985
U67144
--AF315061
AM883187
AF315062
AF168740
AF168746
AY340482
--FJ347645
--AF315038
AF315036
AF315042
ESM Table 2. PCR and sequencing primers
Primer
18SA
18SB
620F
584R
860F
1115R
1324F
1324R
16SarL
16SbrH
OsCO1f
OsCO1r
Sequence 5'-3'
AYCTGGTTGATCCTGCCAGT
ACCTTGTTACGACTTTTACTTCCTC
TAAAGYTGYTGCAGTTAAA
ACGCTATTGGAGCTGGAAT
GAATAATGGAATAGGA
TASGACGGTATCTGATCGTCTT
GGTGGTGCATGGCCG
CGGCCATGCACCACC
CGCCTGTTTATCAAAAACAT
CCGGTCTGAACTCAGATCACGT
AATTATTCGAATTGAATTAGG
AATCAAAATAGGTGTTGGAATAG
References
Medlin et al. (1988)
Nygren and Sundberg (2003)
Nygren and Sundberg (2003)
Persson, pers. comm.
Turbeville et al. (1992)a
Nygren and Sundberg (2003)
Cohen et al. (1998)
Cohen et al. (1998)
Palumbi (1996)
Palumbi (1996)
Glover et al. (2005)
Glover et al. (2005)
--AY436349
FJ347644
FM178480
KF444429
FJ483547
AY646000
ESM Table 3. List of known Osedax. Footnotes: aClade numerals refer to figure 6e, and build upon Vrijenhoek et al (2009). bOTU's based on sequences deposited at GenBank and
names mentioned in the literature using informal taxonomy. cMaximum length of trunk and crown (mm) after preservation. dOTU's are synonyms based on data provided on
GenBank. eDepth estimated from Google Earth using position data reported under GenBank accession number. fBased on data on GenBank, this OTU may be a synonym of 'nude
palp D'.
Cladea
Described species
I
none
I
none
I
none
II
O. antarcticus
II
none
II
none
Undescribed
OTUsb
'green palp'
Depth range
(m)
1820
Type/voucher Sizec
locality
NE Pacific
3
Palps
Pinnules
Roots
Tube
References
red/green
present
robust, lobate
?
Vrijenhoek et al. 2009
'yellow patch'
& 'pinnules'd
'sagami 5'
633–1018
NE Pacific
5
pale
present
?
Vrijenhoek et al. 2009
113e
NW Pacific
?
?
?
?
?
568-650
Antarctica
25
red, striped
absentg
robust, lobate
thin mucous sheath
Pradillon et al. (unpublished, GenBank
FM995539)
This study
'nude palp A'
1820
NE Pacific
25
red
absent
?
?
Jones et al. 2008, Vrijenhoek et al 2009
'nude palp B'
2893
NE Pacific
25
red
absent
?
?
Jones et al. 2008, Vrijenhoek et al 2009
113e-1018
NE Pacific
12
red
absent
?
?
Rouse et al. 2009, Pradillon et al
(unpublished GenBank FM995540)
1018–1820
NE Pacific
12
red
absent
?
?
Vrijenhoek et al. 2009
II
none
II
none
'nude palps' &
'nude palp C'd
& 'sagami 6'd
'nude palp D'
II
none
'nude palp E'
1018
NE Pacific
12
red
absent
robust, lobate
II
none
'nude palp F'
2893
NE Pacific
18
red
absent
robust, lobate
?
Vrijenhoek et al. 2009
II
none
'sagami 8'f
113e
NW Pacific
?
?
?
?
?
III
none
'spiral'
2893
NE Pacific
25
absent
absent
thin filaments
thick? gelatinous
IV
O. frankpressi
1820–2893
NE Pacific
23
red, striped
present
robust, lobate
gelatinous, hemispherical
Pradillon et al (unpublished, GenBank
FM995534)
Braby et al. 2007, Vrijenhoek et al
2009
Rouse et al. 2004
IV
O. mucofloris
30–125
North Sea
15
white to pink
present
robust, lobulate
gelatinous, hemispherical
Glover et al. 2005
IV
O. japonicus
224–250
NW Pacific
12
white to pink
present
robust, lobulate
gelatinous, cylindrical
Fujikura et al. 2006
IV
none
'yellow collar'
385
NE Pacific
18
red
yes
?
Braby et al. 2007
IV
none
'orange collar'
385–1018
NE Pacific
18
red
yes
robust, lobulate
?
Braby et al. 2007
IV
none
'white collar'
& 'sagami 7'd
113e-1018
NE Pacific
6
red, striped
present
robust, lobulate
?
V
O. rubiplumus
1820–2893
NE Pacfic
59
red
present
long, branched
rigid, cylindrical
Vrijenhoek et al. 2009, Pradillon et al.
(unpublished GenBank FM995542)
Rouse et al. 2004
V
O. roseus
'rosy'
633–1820
NE Pacific
24
bright red
present
long, branched
transparent, cylindrical
Rouse et al. 2008
V
none
'sagami 3'
113e
NW Pacfic
?
?
?
?
?
V
none
'sagami 4'
113e
NW Pacfic
?
?
?
?
?
VI?
O. deceptionensis
21
Antarctica
1.2
pale
absent
robust, lobulate
gelatinous, hemispherical
Pradillon et al (unpublished GenBank
FM995537)
Pradillon et al (unpublished GenBank
FM995541)
This report
Vrijenhoek et al. 2009
ESM Figure 1. Frequency distribution of live palp length in Osedax antarcticus sp. nov. based on measurements of the length of the emergent palps in 73 specimens on a region of
bone.
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