Variability in growth rates of long-lived black coral Leiopathes sp. from the Azores (Northeast Atlantic)

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DOI: 10.3354/meps10052
Cite this publication
Five colonies of black coral Leiopathes sp. were collected as bycatch from depths of 293 to 366 m from the Condor, Acor, and Voador seamounts (Azores region). The colonies had axial diameters between 4.9 and 33.1 mm and heights between 43 and 175 cm. Their ages and radial growth rates were estimated using radiocarbon dating. Results indicated that the smallest and largest colonies had similar radial growth rates of 5 to 7 mu m yr(-1), whereas the other 3 colonies had grown more rapidly by a factor of 3 to 5 at similar to 20 to 30 m mu m yr(-1). Colony lifespan ranged between 265 +/- 90 and 2320 +/- 90 yr. Fine-scale sampling along a radial transect from the edge to the center of the 2320 yr old Leiopathes sp. revealed variable growth rates throughout the colony lifespan. Slower radial growth rates of similar to 4 to 5 mu m yr(-1) were recorded over the initial 1600 yr and the last 300 yr of its life span, and a period of more rapid growth (20 mu m yr(-1)) over the intermediate 400 yr of its life. Variability in radial growth rates among colonies resulted in colony ages that were not linearly correlated to colony axis diameter or height. Our findings of great longevity and slow growth rates for Leiopathes sp. agree with other Leiopathes sp. age and growth studies, indicating that colony and population recovery from damage or removal may take centuries to millennia.
Mar Ecol Prog Ser
Vol. 473: 189199, 2013
doi: 10.3354/meps10052
Published January 21
Deep-sea or cold-water coral ecosystems have
have been recognized as important biodiversity hot -
spots in the deep sea (Roberts et al. 2009). The struc-
tural complexity of cold-water corals provides essen-
tial habitat that is used during feeding and spawning
and as nursery grounds for a range of organisms,
including commercially important fish species (Hu -
sebø et al. 2002, Reed 2002, Costello et al. 2005, Me -
taxas & Davis 2005, Roberts et al. 2009). An added
factor that creates a sense of urgency in protecting
these unique organisms is their extreme vulnerability
to disturbance. Several studies have shown that bot-
tom fishing can damage or destroy cold-water corals,
reducing the 3-dimensional complexity of the bottom
topography and leading to decreased faunal bio -
diversity and biomass (Koslow et al. 2001, Hall-
Spencer et al. 2002, Morgan et al. 2005, Althaus et al.
2009, Clark & Rowden 2009).
Despite the realization of damage to these commu-
nities, the recovery of cold-water coral ecosystems
from bottom fishing impacts is poorly known (Al -
thaus et al. 2009). An understanding of cold-water
coral life history traits, such as age, growth rates, and
longevity, is essential to appreciate the nature and
extent of these effects and to evaluate the time scale
of recovery. Cold-water corals can attain ages on the
order of decades to millennia (e.g. Druffel et al. 1995,
Adkins et al. 2004, Andrews et al. 2009, Roark et al.
© Inter-Research 2013 ·*Email:
Variability in growth rates of long-lived black coral
Leiopathes sp. from the Azores
M. Carreiro-Silva
, A. H. Andrews
, A. Braga-Henriques
, V. de Matos
F. M. Porteiro
, R. S. Santos
Centre of IMAR of the University of the Azores, Department of Oceanography and Fisheries/UAz & LARSyS Associated
Laboratory, Rua Prof. Dr Frederico Machado, 4, PT-9901-862 Horta, Azores, Portugal
NOAA Fisheries - Pacific Islands Fisheries Science Center, 99-193 Aiea Heights Drive 417 Aiea, Hawaii 96701, USA
ABSTRACT: Five colonies of black coral Leiopathes sp. were collected as bycatch from depths of
293 to 366 m from the Condor, Açor, and Voador seamounts (Azores region). The colonies had
axial diameters between 4.9 and 33.1 mm and heights between 43 and 175 cm. Their ages and
radial growth rates were estimated using radiocarbon dating. Results indicated that the smallest
and largest colonies had similar radial growth rates of 5 to 7 µm yr
, whereas the other 3 colonies
had grown more rapidly by a factor of 3 to 5 at ~20 to 30 µm yr
. Colony lifespan ranged between
265 ± 90 and 2320 ± 90 yr. Fine-scale sampling along a radial transect from the edge to the center
of the 2320 yr old Leiopathes sp. revealed variable growth rates throughout the colony lifespan.
Slower radial growth rates of ~4 to 5 µm yr
were recorded over the initial 1600 yr and the last
300 yr of its life span, and a period of more rapid growth (20 µm yr
) over the intermediate 400 yr
of its life. Variability in radial growth rates among colonies resulted in colony ages that were not
linearly correlated to colony axis diameter or height. Our findings of great longevity and slow
growth rates for Leiopathes sp. agree with other Leiopathes sp. age and growth studies, indicating
that colony and population recovery from damage or removal may take centuries to millennia.
KEY WORDS: Antipatharia · Cold-water corals · Deep-sea · Growth · Longevity · Radiocarbon ·
Northeast Atlantic
Resale or republication not permitted without written consent of the publisher
Mar Ecol Prog Ser 473: 189199, 2013
2009). Some antipatharians or black corals are partic-
ularly long-lived, with estimated ages in the thou-
sands of years. At the extreme end of longevity is an
estimate exceeding 4000 yr for a Leiopathes sp.
colony in Ha waii (USA), representing the longest-
lived invertebrate to date (Roark et al. 2009).
The Azores harbor diverse cold-water coral garden
communities, dominated by gorgonians, stylasterids,
and black corals, predominantly inhabiting rocky
hard-bottom areas in island slopes and seamounts
(OSPAR Commission 2010a, Braga-Henriques et al.
2012, Tempera et al. 2012). Local artisanal fisheries
have continually targeted some of these coral-rich
areas in the seamounts for several decades, mostly
using bottom-tending longline and handline gears.
The bycatch of bottom longline fisheries revealed
that Leiopathes sp. are often damaged and captured
during fisheries operations (Sampaio
et al. 2012), raising concerns on the
extent of these impairments and their
recovery potential.
The aim of the present study was to
estimate age and growth rates of Leio -
pathes sp. in the Azores region using
radiocarbon dating. Additional objec-
tives were to (1) determine the vari-
ability in growth rates within and
between black coral colonies from
the same and different geographic
regions; and (2) measure the correla-
tion between colony age and skeletal
axis diameter and height as a basis for
bycatch census purposes and video
transect analyses. Ultimately, the goal
is to gather information to better eval-
uate the vulnerability of these species
to fisheries impacts and the capacity of
Leiopathes sp. for recovery from dam-
age or removal.
Specimen collection and identification
Black coral colonies were obtained from bycatch
material from the local longline fishing fleet. Two
specimens were collected from Condor seamount at
depths of 293 and 366 m, 2 other from the Açor
seamount at depths of 302 and 366 m, and a fifth from
the Voador seamount at a depth of 366 m (Fig. 1,
Table 1). The material was deposited in the collection
of the Department of Oceanography and Fisheries of
the University of the Azores (DOP-UAz).
All colonies were identified as belonging to the
genus Leiopathes (Family Leiopathidae) based on
the presence of an irregular sympodial corallum with
multi-directional branching and uniserial to biserial
Specimen Date Site Latitude, longitude Depth Axis diameter Height Morphotype
ID (m) (mm) (cm)
DOP-1985 1985 Açor Bank 38° 17’ N, 28° 52’ W 366 28.9−33.1 175 2
DOP-1356 23 Mar 2007
Açor Bank 38° 17’ N, 28° 52’ W 307 16.5−17.8 155 1
DOP-4588 Jan 2009 Voador Bank 37° 32’ N, 30° 43’ W 366 25−26
137 1
DOP-799 17 Sep 2006 Condor Bank 38° 08’ N, 29° 05’ W 293 11.4−14.2 113 1
DOP-4587 Jan 2009 Condor Bank 38° 08’ N, 29° 05’ W 366 4.9−5.2 43 3
Table 1. Leiopathes sp. Data of the 5 specimens used in this study. Collection year and site (bank) were known for all speci-
mens, with the exception of DOP-1985, where collection year was estimated as 1985 by the fishermen. Axis diameter: mini-
mum and maximum values at the sample extraction plane. Height: vertical colony height.
: Collection year may not be repre-
sentative of the final year of formation for this colony based on observation of encrusting organisms on exterior of basal
: Base axis was at an angle, making a diameter measurement difficult
Fig. 1. Collection sites of Leiopathes sp. in the Condor (DOP-799, DOP-4587),
Açor (DOP-1985, DOP-1356), and Voador (DOP-4588) seamounts, within the
Azores region
Carreiro-Silva et al.: Black coral growth rates
branchlets with a scorpioid arrange-
ment. These branch lets bear small,
simple, and smooth-surfaced spines
reduced in size or absent on larger
branches and stems (Opresko 1998).
Careful examination of some of these
morphological characters (e.g. gross
morphology, size of the colony; length,
diameter, and shape of the terminal
branches, dimension and distribution
of the spines and polyps on the axial
skeleton) suggested that the studied
specimens belonged to 3 morpho-
types: (1) DOP-799, DOP-1356, DOP-
4588, (2) DOP-1985, and (3) DOP-
4587. Because the species currently
classified under the genus Leiopathes
in the NE Atlantic require revision
(D. Opresko pers. comm.), we refrain
from assigning these morphotypes to
specific species. Genetic identification
of these specimens was not possible
due to the lack of freshly preserved
Sample preparation
Cross sections 1 cm thick were
taken from the bases of the colonies
with a hand saw (Fig. 2). The portions
of each colony were examined to
locate the best position for cross sec-
tioning to optimize alignment of the skeletal axis
with the vertical orientation of the micromilling
machine (z-axis). A sample series of skeletal anti -
pathin (a structural protein), oriented along the
radius, was extracted using the micro milling ma -
chine for each colony section. Sample extraction
began at the edge and progressed along radial
lines successively toward the axial core. Each extrac-
tion was made with a 500 µm carbide cutter bit
with a spherical shape (Brasseler USA
, Part No.
H71.11.005). The bit was guided through the skeletal
material at a depth near the bit radius (~250 µm), and
multiple passes were made to obtain enough sample
mass. Near-edge samples were ~250 µm in radial
thickness because of overlapping ex traction paths.
Samples ex tracted closer to the core, and separated
from the successive near-edge samples, had an
extraction path thickness using the full bit diameter
(500 µm). Each extraction was removed successively
with a fine-tipped blade under magnified viewing
conditions. Target extraction mass was near 3 mg at
a minimum for radiocarbon analyses. Because sam-
ples were extracted along radial lines, some extrac-
tion paths were full circle, sampling the entire radius,
to increase sample mass for the period of formation.
In order to obtain enough sample mass, core samples
were typically >500 µm in diameter and deeper than
the near-edge radial extraction.
The selection of sample areas within each colony
depended on the size of the colony and the potential
analysis, either radiocarbon dating or bomb radiocar-
bon dating. Rough estimates of age from growth zone
counting and growth rates from other studies on sim-
ilar corals (e.g. Roark et al. 2006) provided a basis for
the limits of the extraction series (working to encom-
pass pre-bomb to post-bomb Δ
C levels). For most
colonies, the edge was sampled at the finer scale,
stated previously, to evaluate the application of bomb
radiocarbon dating. The locations of older samples
ranging to the core were selected based on the regu-
Fig. 2. Leiopathes sp. (A) Complete type-specimen (height: 62 cm; axis dia -
meter: 18 mm) assigned to morphotype 3 (DOP reference collection). Esti-
mated age is 300 to 1700 yr based on the growth rates provided in this study.
(B) Colony section used in sample extractions for DOP-1985. (C) Frame grab
from micromill computer showing the sample series taken from the edge of
DOP-1985, with an overlap of extraction paths (0.5 mm bit width). (D) Post-ex-
traction surface of the coral section showing the narrower sample series at the
edge, 3 intermediate radial samples (full bit width at 0.5 mm), and the axial
core extraction
Mar Ecol Prog Ser 473: 189199, 2013
lar spacing and avoidance of fractures or separations
in the skeletal structure. Pre-bomb samples from the
fine scale series near the edge (determined later) and
the older samples ranging to the core were used in
radiocarbon dating. Extraction of coral material from
the 5 coral colonies successfully produced >50 sam-
ples to provide a selection of measurement locations
in each colony section. Samples were analyzed in
successive sets, the results of which were used to
make educated decisions for the next set of samples.
Ultimately, 25 samples were selected and analyzed
with sample mass ranging from 1.0 to 7.3 mg and the
number per colony ranging from 1 in the smallest
colony to 10 in the largest colony (Table 2).
Analytical measurements
Extracted samples were submitted as powder to the
Center for Accelerator Mass Spectrometry (CAMS) at
Lawrence Livermore National Laboratories, Liver-
more, California, USA, for radiocarbon analysis. Sam-
ple CO
was generated via combustion, converted to
graphite and measured for radiocarbon (
C) content
using an accelerator mass spectrometer (AMS). The
C values were reported as Δ
C (Stuiver & Polach
1977) and were corrected for isotopic fractionation us-
ing measured δ
C values and an assumed δ
C value
of −17‰, rounded off from a mean of the measured
values (n = 9, 17.4 ± 0.8‰ SD). For bomb radiocarbon
dating, the Δ
C measured in the latest samples were
used to determine rough estimates of age and growth
from post-bomb to pre-bomb levels. For radiocarbon
dating, the full series of pre-bomb Δ
C values were
used to determine colony age, and were reported as
conventional radiocarbon age (CRA) and calendar
year before present (BP, relative to 1950 AD) corrected
reservoir effects (ΔR). A calculated reservoir age of
399 ± 17
C yr was determined from a mean Δ
value of −49 ± 2‰ (n = 9), reported for 1955 from the
North Atlantic Ocean (Stuiver 1980), which was con-
verted to the fraction modern (Fm) for 1950 (Fm =
0.9516), and then
C years with the following conver-
sion (
C yr = −8033 × ln(Fm)). A global marine reser-
voir correction age of 464 ± 23
C yr (from the Mar-
ine09 supplementary material in Reimer et al. 2009)
Specimen ID CAMS ID Sample ID δ
C (‰) Fm ± 1 SD Δ
C ± 1 SD (‰) CRA ± 1 SD (yr)
DOP-1985 147817 AA 09 −17 0.9549 ± 0.0035 −52.0 ± 3.5 370 ± 30
147381 AA 10 −17 0.9509 ± 0.0049 −55.9 ± 4.9 405 ± 45
147382 AA 11 −16.6 0.9357 ± 0.0028 −71.1 ± 2.8 535 ± 25
147383 AA 12 −17 0.9330 ± 0.0029 −73.8 ± 2.9 555 ± 30
147384 AA 14 −17 0.9299 ± 0.0031 −76.8 ± 3.1 585 ± 30
147385 AA 15 −17 0.9241 ± 0.0029 −82.6 ± 2.9 635 ± 25
147818 AA 17 −17 0.9049 ± 0.0040 −101.6 ± 4.0 800 ± 40
147819 AA 18 −16.4 0.8705 ± 0.0036 −135.8 ± 3.6 1115 ± 35
147820 AA 19 −17 0.7862 ± 0.0030 −219.5 ± 3.0 1935 ± 35
147386 AA 20 −17 0.7283 ± 0.0025 −277.0 ± 2.5 2545 ± 30
DOP-1356 147824 AA 44 −17.6 0.9520 ± 0.0028 −54.9 ± 2.8 395 ± 25
147825 AA 46 −17 0.9445 ± 0.0033 −62.4 ± 3.3 460 ± 30
147829 AA 45 −17.4 0.9387 ± 0.0034 −68.1 ± 3.4 510 ± 30
147830 AA 47 −17.2 0.9354 ± 0.0027 −71.3 ± 2.7 535 ± 25
147823 AA 43 −17 0.9259 ± 0.0030 −80.8 ± 3.0 620 ± 30
DOP-4588 147826 AA 21+22
−18.9 1.0945 ± 0.0036 86.6 ± 3.6 >Modern
147831 AA 23 −17 1.1006 ± 0.0034 92.7 ± 3.4 >Modern
147821 AA 33 −16.6 0.9303 ± 0.0039 −76.4 ± 3.9 580 ± 35
147827 AA 34 −18.1 0.9196 ± 0.0028 −87.0 ± 2.8 675 ± 25
147387 AA 35 −17 0.9046 ± 0.0028 −102.0 ± 2.8 805 ± 30
DOP-799 147388 AA 37 −17 1.1040 ± 0.0142 96.0 ± 14.2 >Modern
147828 AA 40 −18.2 0.9562 ± 0.0033 −50.7 ± 3.3 360 ± 30
147822 AA 36 −17 0.9459 ± 0.0035 −61.0 ± 3.5 445 ± 35
147389 AA 39 −17 0.9372 ± 0.0030 −69.5 ± 3.0 520 ± 30
DOP-4587 147390 AA 42 −17 0.9323 ± 0.0041 −74.4 ± 4.1 565 ± 40
Table 2. Leiopathes sp. Results from radiocarbon assays separated by colony (Specimen ID) and into the consecutive sample
series taken along each radial axis (CAMS ID = Center for Accelerator Mass Spectrometry lab number and Sample ID = labo-
ratory record). Provided are assumed (−17‰) and measured (given to 0.1‰) δ
C values, with the measured radiocarbon frac-
tion modern (Fm), calculated Δ
C, and conventional radiocarbon age (CRA). >Modern: >1950 AD.
: Samples combined to
increase sample extraction mass
Carreiro-Silva et al.: Black coral growth rates
was subtracted from the reservoir age (399 ± 17
yr) for a ΔR value of −65 ± 29
C yr. The negative re-
sult is consistent with more centrally located ocean
positions, such as Hawaii (ΔR = −28 ± 4
C yr; Roark
et al. 2006; cf. coastal regions affected by upwelling
with positive ΔR values, e.g. Sherwood et al. 2008,
Soares & Martins 2010). Calibrated calendar age was
determined using the calculated ΔR value in Calib 6.0
(http:// with the Marine09 calibration
record (Reimer et al. 2009). Because the sample ex-
traction was composed of material formed over
several decades of deposition, the Calib 6.0 calculation
option of a ‘sample calendar year span’ was chosen in
the Calib 6.0 analysis (Stuiver & Reimer 1993). This
option performs a moving average of the calibration
curve and is the best option for this kind of sample de-
sign, and the final reported age for each sample was
the median probability age. Final sample age deter-
minations were corrected to the time of collection by
adding 35 to 59 yr, depending on the sample. Based
on these age data, long-term average growth rates for
full colonies and within colonies were determined and
plotted for interpretation.
Radiocarbon age
Fm measured for
C ranged, as expected, from a
low ratio in the core of the largest colony to >1.0
(>Modern) for the most recent and post-bomb affected
samples near or at the edge of 2 of the 5 colonies
(Table 2). Calculated Δ
C values ranged from a low of
−277.0 ± 2.5‰ (mean ± SD) from the core of DOP-
1985 to post-bomb elevated values approaching
100‰ in the near-edge material of DOP-799 and
DOP-4587. Change in Δ
C across the radial transects
of each colony revealed a variable rate of decline with
3 of the 5 colonies having similar rates of decline
(Fig. 3). This corresponded to conventional radiocar-
bon ages ranging from near 500 to 600
C yr for the
smallest colonies to >2000
C yr for the largest colony.
The age of individual extracted samples and total
colony age (lifespan of the colony) were determined
using the calibrated radiocarbon ages (Table 3). Only
measured Δ
C values less than the value used to
determine a ΔR could be used in determining a cal-
endar year before 1950; hence, some pre-bomb val-
ues were deemed ‘Modern’ and likely formed close
to 1950, but were not used in the determination of
sample age. The age of the 2 smallest colonies was
near 300 yr, and the oldest was ~2320 yr old. Dis-
tances to the center of the extraction path (cut width)
were plotted with radiocarbon age to reveal differ-
ences and similarities in growth rates over time, both
within and between colonies (Fig. 4).
Growth rates
Growth rates differed among the Leiopathes sp.
specimens studied. The smallest and largest colonies
had similar growth rates of 5 to 7 µm yr
, whereas the
other 3 colonies grew more rapidly by 3 to 5 times,
with growth rates of ~20 to 30 µm yr
(Table 4). The
largest colony was the slowest-growing overall, but
the within-colony rates differed over time (Fig. 5).
The earliest and latest growth rates were similar at 4
Radial distance from ed
e (mm)
C (‰)
12 14 16
Fig. 3. Leiopathes sp. Radial distance from axis to outer edge
(mm) versus Δ
C determinations for the 5 colonies studied
Radial distance from edge (mm)
C age (cal yr BP)
Fig. 4. Leiopathes sp. Radial distance from axis to outer edge
versus reservoir-corrected calendar age for the 5 colonies
Mar Ecol Prog Ser 473: 189199, 2013
to 5 µm yr
, with a period of
more rapid growth at ~20 µm yr
through the upper middle ages of
the colony (2 to 8 mm in radius
and ~400 to 750 yr ago). In addi-
tion, growth rates of the other 3
colonies were consistent with the
faster growth period of the
largest colony. Colony ages
across the 5 specimens used in
this study did not significantly
correlate with the main axis di-
ameter of the colony (r = 0.76, p =
0.14, n = 5) or their height (r =
0.59, p = 0.29, n = 5), meaning
that axis diameter and colony
height did not increase linearly
with colony age based on our lim-
ited number of samples.
Use of bomb radiocarbon dating
provided estimates of age and
growth that support the conven-
tional radiocarbon dating, but the
findings were not well defined.
Because of low growth rates and
the relatively short time frame for
bomb radiocarbon dating (~50 yr),
there was not enough material for
good temporal resolution. Based
on changes in Δ
C from what can
be classified as pre-bomb to post-
bomb values near the edge of the
coral section, 2 colonies provided
estimates of age. For DOP-799,
the assumption was made that
sample extraction was before and
near the initial rise of bomb Δ
C and a limit of calen-
dar year 1955 was chosen. Support for this determina-
tion comes from the measured value of −50.7 ± 3.3‰,
similar to the value (mean ± SD) determined for the
North Atlantic Ocean pre-bomb levels (−49 ± 2‰;
Stuiver 1980). A conservative growth rate estimate
based on the extremes of extraction radius provided a
rate of ~10 to 20 µm yr
. For DOP-4588, it was as-
sumed that the second sample measurement (92.7 ±
3.4‰) encompassed the rise and peak for bomb Δ
Given that the sample crossed the peak years, 1965
was arbitrarily chosen as the year of formation. A con-
servative growth rate estimate was approximately 10
to 25 µm yr
. Bomb radiocarbon dating estimates for
2 of the other 3 colonies was not possible be cause ele-
vated levels were not measured due to other factors
(edge not sampled or no post-bomb growth).
Specimen Sample Distance Cut width Calendar Age
ID ID (mm) (mm) years BP (yr)
DOP-1985 AA 09 0.15 0.30 Modern Modern
AA 10 0.43 0.26 95 (5−135) 130 (40−170)
AA 11 0.69 0.26 245 (195−295) 280 (230−330)
AA 12 0.95 0.26 265 (215−320) 300 (250−355)
AA 14 1.20 0.26 300 (255−350) 335 (290−385)
AA 15 1.46 0.26 345 (300−390) 380 (330−425)
AA 17 4.25 0.50 490 (450−525) 525 (485−560)
AA 18 8.25 0.50 720 (675−765) 755 (710−800)
AA 19 12.35 0.50 1560 (1500−1620) 1600 (1540−1660)
AA 20 16.00 0.30 2290 (2230−2340) 2320 (2270−2380)
DOP-1356 AA 44 0.15 0.30 74 (0−100) 130 (60−160)
AA 46 1.83 0.50 160 (105−240) 215 (160−300)
AA 45 4.68 0.50 205 (180−270) 265 (235−325)
AA 47 5.98 0.50 245 (195−290) 300 (250−350)
AA 43 8.23 0.30 340 (285−380) 395 (340−440)
DOP-4588 AA 21+22 0.28 0.28 >Modern
AA 23 0.70 0.14 >Modern
AA 33 5.95 0.50 295 (250−355) 355 (305−415)
AA 34 9.75 0.50 380 (330−430) 440 (385−485)
AA 35 13.03 0.30 490 (465−520) 550 (520−580)
DOP-799 AA 37 0.15 0.30 >Modern
AA 40 0.95 0.50 Modern
AA 36 2.90 0.50 135 (75−230) 195 (130−285)
AA 39 7.12 0.26 220 (185−275) 275 (240−335)
DOP-4587 AA 42 1.75 0.50 275 (220−340) 335 (280−395)
Table 3. Leiopathes sp. Synopsis of radial samples extracted from the coral axis, given
as the distance from the section edge toward the center (distance to the center of extrac-
tion width for mean growth zone radius), with the width of the cut from milling based on
the bit diameter and extraction (successive radial path or single path using full tip diam-
eter). Listed are the calculated calendar years before present (BP; median probability
age) with the uncertainty from Calib 6.0 (moving average) with a re servoir age (ΔR) of
−65 ± 29 yr (ca. Δ
C = −49 ± 2; Stuiver 1980; mean ± SD). Age was corrected to time of
collection from the calibration year of 1950. Data in parentheses are 1 SD calibrated age
ranges. >Modern: >1950 AD.
: Δ
C value too close to the pre-bomb reference used in
determining ΔR, which leads to an error in Calib 6.0 age calculations
0 2 4 6 8 1012141618
Radial distance from edge (mm)
C age (cal yr BP)
Slope = 0.0050 mm yr
Slope = 0.018 mm yr
Slope = 0.0049 mm yr
Fig. 5. Leiopathes sp. Radial distance from axis to outer edge
versus reservoir-corrected calendar age for the largest colony
studied (DOP-1985), with variable growth rates
Carreiro-Silva et al.: Black coral growth rates
Radiocarbon dating
Radiocarbon dating indicated that some Leio pathes
sp. black corals in the Azores have been continuously
growing for at least 2300 yr with growth rates as low
as 5 µm yr
. These findings corroborate previous
reports of high longevity and slow growth rates for
Leiopathes sp. in the Pacific and NW Atlantic (Roark
et al. 2006, 2009, Williams at al. 2006, Prouty et al.
2011), demonstrating that these life history traits are
consistent across oceanographic regions of the world.
Our age estimates for Leiopathes sp. fall within the
lower to middle range of reported values for other
Leiopathes sp. studies (198 to 4265 yr: Roark et al.
2006, 2009, Williams et al. 2006, Prouty et al. 2011).
However, the maximum age reported here was not
from the tallest and thickest colony recorded from
the region, suggesting that Leiopathes sp. in the
Azores may attain ages exceeding this estimate. A
large specimen that was collected from Condor
Seamount had a height of 3 m and an axis diameter of
40 mm. Unfortunately, this specimen belonged to a
private collection and we were unable to sample it for
radiocarbon dating. However, 1 of the goals of this
study was to estimate age for colonies in the field
from axis diameter. Although correlation between
age estimates and axis diameter resulted in a wide
range of uncertainty, an age interval for this speci-
men was calculated using the range of growth rates
of the largest colony aged in this study (DOP-1985;
6.6 to 7.2 µm yr
radial growth rate). Assuming the
observed changes in growth rate over the lifespan
would be similar for a colony of this size, an estimate
of 2800 to 3000 yr was determined. Given the full
range of measured radial growth rates in this study
(5.2 to 26.0 µm yr
), age could be between 700 and
3900 yr. However, it is likely that a mean growth rate
is most applicable to a colony this large and an age
near 3000 yr is likely. According to this estimation the
theoretical maximum age of Leiopathes sp. in the
Azores is similar to the maximum age reported for
Leiopathes sp. in Hawaii.
Growth rates of our samples were comparable to
those reported for the Gulf of Mexico (8 to 22 µm yr
Prouty et al. 2011), but greater than those given for
Hawaii (2 to 13 µm yr
: Roark et al. 2006, 2009). The
reasons for these differences are uncertain, but may
be related to factors such as food availability dis-
cussed in more detail in the next section. Despite
these differences, growth rates in Leiopathes sp. are
generally lower and lifespans greater than in other
deep-sea coral species. For example, the deep-sea
antipatharian species Stauropathes arctica off New-
foundland (Canada) grows faster and has a shorter
lifespan than Leiopathes sp. (growth rate = 33 to
66 µm yr
, maximum reported lifespan = 82 yr: Sher-
wood & Edinger 2009). Octocorals in general have
more rapid growth rates and lower lifespans than
Leiopathes sp., with growth rates between 50 and
440 µm yr
and an estimated longevity in hundreds
of years (Andrews et al. 2002, Sherwood et al. 2006,
Watling et al. 2011). The only other species with com-
parable growth and longevity to Leiopathes sp. is the
zo anthid Gerardia sp. (lifespan in the order of millen-
nia with growth rates as low as 5 µm yr
: Druffel et
al. 1995, Parrish & Roark 2009, Roark et al. 2009).
Variability in growth rates
We estimated age and growth rates of 5 Leiopathes
sp. colonies of different sizes to explore whether vari-
ations in age and growth were related to colony size.
Growth rates differed among the colonies, with the
largest and smallest colonies presenting the slowest
mean growth rates (5 to 7 µm yr
), and 3 intermedi-
ate sized colonies presenting 4 to 5 times higher
mean growth rates (21 to 26 µm yr
). This may be
related to differences in colony size, colony collection
site, or species-specific differences.
Specimen Maximum age Mean radius Radius range Mean growth rate Growth rate range
ID (yr) (mm) (mm) (mm yr
) (mm yr
DOP-1985 2320 (2270−2380) 16.00 15.60−16.40 0.0069 0.0066−0.0072
DOP-1356 395 (340−440) 8.23 7.95−8.50 0.021
DOP-4588 550 (520−580) 13.03 12.97−13.07 0.024 0.022−0.025
DOP-799 275 (240−335) 7.12 6.17−8.07 0.026 0.018−0.033
DOP-4587 335 (280−395) 1.75 1.55−1.95 0.0052 0.0039−0.0070
Table 4. Leiopathes sp. Mean colony growth rates based on the maximum age estimate. Mean radius and radius range were de-
termined to account for radial growth variation.
: Estimated last year of formation at 1950 leads to a calculated growth rate of
0.024 (0.021−0.030) mm yr
(consideration for evidence that no post-bomb growth occurred)
Mar Ecol Prog Ser 473: 189199, 2013
The morphological characters of the coral colonies
indicated 3 distinct morphotypes. The largest, the
smallest, and the 3 intermediate sized colonies repre-
sented 1 morphotype each. Because the species clas-
sified under the genus Leiopathes in the NE Atlantic
require revision (D. Opresko pers. comm.), and we
did not have enough freshly preserved material for
genetic identification, we were unable to determine
whether the 3 morphotypes were different species.
Recent molecular phylogenetic studies on deep-sea
bamboo corals have demonstrated high lability in
morphological characters traditionally used for their
classification (Dueñas & Sanchez 2009) i.e. deep-sea
corals may show considerable phenotypic plasticity
within species.
If the 3 morphotypes correspond to different spe-
cies, then the observed differences in growth rates
may correspond to species-specific rates. Consider-
able growth rate differences have been reported for
closely related antipatharian species of the genus
Antipathes (Roark et al. 2006, Love at al. 2007).
Growth rates of A. dichotoma from 50 m depth off
Hawaii (Roark et al. 2006) were up to 8 times greater
than growth rates reported for A. dendrochristos col-
lected at 106 m depth off southern California (Love et
al. 2007).
Alternatively, different growth rates may reflect
differences in food availability around the seamount.
The differences in growth rates were recorded for
specimens collected within the Açor (DOP-1985,
DOP-1356) and Condor Seamounts (DOP-799, DOP-
4587, Table 4). Isotopic evidence indicates that anti -
patharian corals feed on suspended matter derived
from surface production (Roark et al. 2006, 2009,
Sherwood et al. 2008), which can be influenced by
local hydrographic patterns. In the Azores, informa-
tion on near-bottom oceanographic conditions (cur-
rents, temperatures, and biogeochemical parame-
ters) is scarce and regional models provide only
coarse information; however, there is evidence that
current circulation and local upwelling− downwelling
patterns around seamounts are distinct from the sur-
rounding ocean in the Northeast Atlantic Ocean, pro-
ducing a diverse and complex physical environment
(Vilas et al. 2009, Mendonça et al. 2012). Preliminary
data on particulate organic carbon (POC) influx for
Condor Seamount suggest that POC concentrations
can be nearly 2 orders of magnitude greater on
the northern slope relative to the southern slope
(A. Colaço pers. comm.). This spatial heterogeneity
in particle flux at the microhabitat scale could ex -
plain differences observed in coral growth rates on a
seamount. Large intra-seamount variability in POC
concentrations and plankton biomass has also been
recorded at the Sedlo and Seine Seamounts located
in subtropical waters of the northeast Atlantic Ocean,
related to differences in local circulation around the
seamount (Vilas et al. 2009).
Growth rate comparisons of Leiopathes from differ-
ent oceanographic regions suggest that surface-layer
productivity, and thus food availability, may influ-
ence coral growth rates. We measured, rates similar
to those reported for the Gulf of Mexico, and greater
than those calculated for Hawaii. Primary productiv-
ity estimates in the Azores region (385 mg C m
are similar to the Gulf of Mexico (417 mg C m
but are nearly 2 times greater than for the Hawaii
region (228 mg C m
, www. seaaroundus. org),
reflecting similarities in the trends observed for Leio -
pathes growth rates among these regions. Regional
differences in growth rates of bamboo coral in New
Zealand have also been attributed to differences
in oceanographic conditions and food availability
(Tracey et al. 2007).
A third potential explanation relates to variations
in growth rates throughout the life span of the coral,
rather than to species-specific growth rates. Roark et
al. (2006) found greater radial growth rates over the
initial 400 yr of a 2370 yr old Leiopathes sp. from
Hawaii. More recently, Prouty et al. (2011) reported
an inverse relationship between colony age and
growth rates based on measurements in 5 colonies
with different axis diameters. Our results show a con-
trasting pattern, with lowest growth rates (4 to 5 µm
) recorded over the initial 1600 yr and the last 300
yr of lifespan of the 2320 yr old Leiopathes sp., with a
period of more rapid growth (20 µm yr
) during the
intermediate 400 yr of life. Similar growth rates were
recorded for our smallest specimen with a lifespan of
300 yr. These results suggest low growth rates in
young and old colonies. Low initial growth rates
could be related to low surface area exposed to the
currents delivering organic particles on which the
polyps are feeding, thus capturing only few re -
sources for growth, as suggested for bamboo corals
from the Gulf of Alaska (Roark et al. 2005). In this
scenario, the growth rate would increase as the coral
surface area and feeding efficiency increased, but
may slow down as the colony reaches a maximum
size. Studies on the growth and modularity of colo-
nial cnidarians suggest that growth rates and colony
maximum sizes are controlled by size-dependent
interactions between the colony and the environ-
ment, such as the balance between metabolic rate
and resource capture (e.g. Sebens 1982, Kim &
Lasker 1998). As the size of the colony increases, the
Carreiro-Silva et al.: Black coral growth rates
amount of resources captured by different parts of
the colony are reduced by colony self-shading. In
addition, energy allocation to reproduction and other
physiological functions may be favored in relation to
growth (Kim & Lasker 1998).
An additional consideration with the observed
growth rate variability is the reliance of radiocarbon
dating on the stability of the
C reference record
over time. It is important to note that a series of
assumptions are necessary in this instance because
time-specific radiocarbon records, extending back
2000 yr, do not exist as a reference in the region. It is
assumed that the calculated ΔR value for the mixed
layer is relatively consistent over time, but minor
variations due to upwelled dissolved inorganic car-
bon or POC (depending on the primary source for
skeletal growth) could change the reservoir age for
unknown periods of time. A minor depression of Δ
on the order of 10‰ can lead to a positive change in
reservoir age of 85
C yr for this study. This factor
was described for the northeast Atlantic from histori-
cal samples, primarily from the ocean margin, and
was attributed to the North Atlantic Oscillation (Tis-
nérat-Laborde et al. 2010). A problem with age esti-
mates determined in this study is unlikely because
(1) sample extraction covered decades of growth
and likely normalizes such minor variations (>10 yr
period) and (2) the Azores are more closely associ-
ated with the structural stability of the North Atlantic
Gyre; nevertheless, it is important to mention this
potential complication when making age determina-
tions on deep-sea corals.
Management and conservation
As is the case with old-growth forests of terrestrial
environments, our findings indicate that Leiopathes
are particularly vulnerable to natural and anthro-
pogenic disturbance. Once removed, these colonial
organisms will not return to their former structural
and ecological significance within our lifetime. Dis-
turbance and removal from bottom trawling is among
the most significant threats to cold-water corals
(Althaus et al. 2009), and recent studies have demon-
strated that other bottom-set fishing gears, such as
longlines, can also significantly impact these commu-
nities in areas of intense fishing (Duran Muñoz et al.
2011, Sampaio et al. 2012). In the Azores, bottom
trawling and deep-sea netting are forbidden and
only bottom long lining is allowed (European Council
Regulation [EC] No. 1568/2005 of 20 September
2005; see also Probert et al. 2007, Santos et al. 2009).
Recent studies on bycatch during longline fishing
experiments revealed that 33% was composed of
corals, and ~7% of the corals caught were anti pa tha -
ri ans (DOP-UAz unpublished data).
Anecdotal reports from fishermen suggest a pro-
gressive reduction in the accidental capture of
Leiopathes after 20 yr of bottom-longline fishing on
insular slopes and seamounts. Large Leiopathes
colonies (>2 m tall), and previously collected by fish-
ermen, were not recorded in video surveys recently
conducted in the Azores (A. Braga-Henriques pers.
obs.). This finding could indicate that the largest and
oldest colonies have already been removed from the
population in heavily fished areas. These observa-
tions suggest that bottom longlines have had a de -
structive effect, and management strategies to
reduce impacts to these ancient corals should be
The Government of the Azores recently created 4
deep-sea marine protected areas for the conservation
of particularly sensitive benthic habitats, such as
coral and sponge reefs and gardens. These protected
areas are the Sedlo Seamount (Santos et al. 2009,
2010), and 3 high seas areas claimed by Portugal to
the Commission on the Limits of the Continental
Shelf (Altair and Antialtair Seamounts, and an area
of the Mid-Atlantic Ridge north of the Azores), of
which all are part of the OSPAR network of Marine
Protected Areas (OSPAR Commission 2010b). The
present study on Leiopathes contributes to this con-
servation effort by providing estimates of growth and
longevity for an important structural member of
these habitats, one of the longest-lived corals known.
This information must be considered for the classifi-
cation of additional vulnerable habitats that harbor
ancient communities and deserve protection.
Acknowledgements. IMAR-DOP/UAz (Research and Devel-
opment Unit no. 531) and LarSyS-Associated Laboratory are
supported by the Portuguese Foundation for Science and
Technology (FCT) under a strategic project and DRCTC-GR
Azores through a Pluriannual Funding scheme. This study
was supported by the EU-funded CoralFISH project, Marie
Curie International Reintegration Grant (to M.C.-S.,
PIRG03-GA-2008-231109), post-doctoral fellowship from
FCT (to M.C.-S., SFRH-BPD-34634-2007), and a PhD schol-
arship from Direcção Regional para a Ciência e a Tecnologia
(to A.B.-H., M3.1.2/F/016/2008). The service charges for this
open access publication have been covered by LARSyS
Associated Laboratory through FCT/MCE project PEst-OE/
EEI/LA0009/2011. We thank bottom long-line fishermen, in
particular J. Gonçalves, for providing specimens. P. Reimer
(Queen’s University, Belfast), E. Druffel (University of Cali-
fornia, Irvine), O. Sherwood (Dolan Integration Group), and
T. Brown (Lawrence Livermore National Laboratory) are
Mar Ecol Prog Ser 473: 189199, 2013
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was performed by T. Brown at the Center for Accelerator
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Medeiros at DOP-UAc. We thank F. Parrish and R. Nichols
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Editorial responsibility: Karen Miller,
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Submitted: December 6, 2011; Accepted: September 17, 2012
Proofs received from author(s): December 28, 2012
  • ... They create a complex three-dimensional habitat and support high levels of biodiversity, providing refuge, feeding opportunities, and spawning and nursery areas for a wide range of organisms (Buhl-Mortensen et al., 2010). Cold-water corals grow extremely slowly (a few to several mm per year) and can live for hundreds or thousands of years (e.g., Roberts et al., 2009;Watling et al., 2011;Carreiro-Silva et al., 2013). The limited knowledge on the distribution and extent of cold-water coral habitats makes it difficult to assess changes. ...
    Full-text available
    To understand the restoration potential of degraded habitats, it is important to know the key processes and habitat features that allow for recovery after disturbance. As part of the EU (Horizon 2020) funded MERCES project, a group of European experts compiled and assessed current knowledge, from both past and ongoing restoration efforts, within the Mediterranean Sea, the Baltic Sea, and the North-East Atlantic Ocean. The aim was to provide an expert judgement of how different habitat features could impact restoration success and enhance the recovery of marine habitats. A set of biological and ecological features (i.e. life-history traits, population connectivity, spatial distribution, structural complexity and the potential for regime shifts) were identified and scored according to their contribution to the successful accomplishment of habitat restoration for five habitats: seagrass meadows, kelp forests, Cystoseira macroalgal beds, coralligenous assemblages and cold-water coral habitats. The expert group concluded that most of the kelp forests features facilitate successful restoration, while the features for the coralligenous assemblages and the cold-water coral habitat did not promote successful restoration. For the other habitats the conclusions were much more variable. The lack of knowledge on the relationship between acting pressures and resulting changes in the ecological state of habitats is a major challenge for implementing restoration actions. This paper provides an overview of essential features that can affect restoration success in marine habitats of key importance for valuable ecosystem services
  • ... This might be because small colonies could not be seen on the videos as they are very thin and flattened against the substrate. Antipatharians are amongst the oldest known marine organisms, with very slow growth and millennial longevities (Roark et al., 2009;Carreiro-Silva et al., 2013). Leiopathes glaberrima and accompanying benthic invertebrates have been reported as Vulnerable Marine Ecosystem indicators and protection measures should be adopted as a precautionary approach (Massi et al., 2018). ...
  • ... The species belonging to this genus are characterised by an extraordinary longevity, being considered among the oldest-living creatures on Earth (Roark et al. 2006). The radiocarbon dating of a Leiopathes colony from Hawaiian waters revaled an extraordinary age of 4265 years (Roark et al. 2006;Wagner and Opresko 2015) and Atlanto-Mediterranean L. glaberrima colonies were found to be as old as 2000 years (Carreiro-Silva et al. 2013;Bo et al. 2015). Although growth rates vary among individuals and depends on locality and depth, the occurrence of millennial specimens was hypothesised to reflect a long-term stability of undisturbed deep biocoenoses, particularly surprising for a semi-enclosed, heavily exploited basin such as the Mediterranean Sea (Bo et al. 2015). ...
    The term cold-water coral sensu lato groups taxa with a more or less pronounced frame-building ability (e.g. Lophelia pertusa and Madrepora oculata) with forest-forming organisms both on hard (e.g. Leiopathes glaberrima, Parantipathes larix, Callogorgia verticillata and Viminella flagellum) and soft bottoms (e.g. Isidella elongata, Funiculina quadrangularis and Kophobelemnon stelliferum). Cold-water coral species and their occurrence in the Mediterranean Sea are here reviewed and discussed from a biogeographic point of view, considering geographical areas of occurrence and bathymetric distribution. The present-day occurrence of living cold-water corals is then compared to the main deep currents of the Mediterranean Sea. Due to the proper interaction between topography and a combination of cold, oxygenated and trophic-carrying water masses (i.e. Levantine Intermediate Water, deep waters and cascading effects), cold-water coral communities develop in a mosaic-like situation along the main paths that such water masses follow within the basin. Finally, knowledge gaps and future perspectives in the study of cold-water coral occurrence, distribution and biogeography are highlighted. The currently still scarce knowledge on the Mediterranean deep-sea and on the basin-scale distribution of the most important cold-water corals species represents crucial biogeographical information. This gives fundamental indications on the location of the Mediterranean vulnerable deep marine ecosystems for future management strategies.
  • ... Furthermore, the information that is available is typically qualitative in nature and tends to be based on expert opinion relating to the spatial distribution of activities/pressures and the restorative aspects of the habitat's key species. For example, although little information is available on the recovery potential of deep-sea features, there is a general consensus that highly impacted corals are unlikely to recover at relatively short-medium temporal scales due to their slow growth rate, high longevity, long reproductive cycles and low rates of recruitment coupled with the continuously increasing degree of human-induced impacts [38][39][40][41]. ...
  • ... Many of the species that live on seamounts are slow-growing, long-lived and slow to reproduce. Some cold-water corals are known to live for hundreds to thousands of years (Roark et al. 2006;Rogers et al. 2007;Carreiro-Silva et al. 2013). Schlacher et al. (2014) considered that the life history characteristics of the cobalt crust fauna would make recovery from the mechanical impacts of mining very slow. ...
    The new industry of deep-sea mining (DSM) potentially offers abundant supplies of several metals from the deep ocean, but the ores will need to be recovered from pristine environments in which the ecosystems are often poorly known. Information that is available for some of these environments suggests that organisms may struggle to recover from the impacts of DSM, whilst in other areas the impacts may be somewhat less.
  • ... Leiopathes sp. has a slow radial growth rate of 0.005-0.022 mm yr −1 and is one of the longest living species on earth (Roark et al., 2009;Carreiro-Silva et al., 2013). The slower radial growth rates and their longevity might be of an advantage in areas where food supply is scarcer. ...
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    Cold-water coral carbonate mounds, created by framework-building scleractinian corals, are also important habitats for non-scleractinian corals, whose ecology and role are understudied in deep-sea environments. This paper describes the diversity, ecology and role of non-scleractinian corals on scleractinian cold-water coral carbonate mounds in the Logachev Mound Province, Rockall Bank, NE Atlantic. In total ten non-scleractinian species were identified, which were mapped out along eight ROV video transects. Eight species were identified as black corals (three belonging to the family Schizopathidae, one each to the Leiopathidae, Cladopathidae, and Antipathidae and two to an unknown family) and two as gorgonians (Isididae and Plexauridae). The most abundant species were Leiopathes sp. and Parantipathes sp. 2. Areas with a high diversity of non-scleractinian corals are interpreted to offer sufficient food, weak inter-species competition and the presence of heterogeneous and hard settlement substrates. A difference in the density and occurrence of small vs. large colonies of Leiopathes sp. was also observed, which is likely related to a difference in the stability of the substrate they choose for settlement. Non-scleractinian corals, especially black corals, are an important habitat for crabs, crinoids, and shrimps in the Logachev Mound Province. The carrier crab Paromola sp. was observed carrying the plexaurid Paramuricea sp. and a black coral species belonging to the genus Parantipathes, a behavior believed to provide the crab with camouflage or potentially a defense mechanism against predators. More information on the ecophysiology of non-scleractinian corals and fine-scale local organic matter supply are needed to understand what drives differences in their spatial distribution and community structure.
  • ... Deep-sea species are inherently vulnerable to environmental change. Characteristics of deep-sea organisms include increased longevity, slow growth rates, reproduction late in life and low fecundity (Carreiro-Silva et al., 2013;Levin et al., 2016;Danovaro et al., 2017b;Montero-Serra et al., 2018). These particular life history strategies mean that many deep-sea species have an increased sensitivity to human activities such as mining, fisheries and climate change. ...
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    Commercial seabed mining seems imminent, highlighting the urgent need for coherent, effective policy to safeguard the marine environment. Reconciling seabed mining with the United Nations Sustainable Development Goals will be difficult because minerals extraction will have irreversible consequences that could lead to the loss of habitats, species and ecosystems services. A dialog needs to take place around social, cultural, environmental and economic costs and benefits. Governance of human interactions with the seabed is fragmented and lacks transparency, with a heavy focus on facilitating exploitation rather than ensuring protection. In the light of high uncertainties and high stakes, we present a critical review of proposed policy options for the regulation of seabed mining activities, recommend actions to improve seabed governance and outline the alternatives to mining fragile deep-sea ecosystems.
  • Chapter
    Antipatharians are characterized by an erect, rigid chitinous skeleton that create long unbranched whip-like coil or tree-like, unbranched colony. The skeleton of black corals represents a structure typical for a laminated composite. However, the detailed inorganic-organic composition can differ from one species to another. Different elements that can be found in the skeleton are carbohydrates, lipids, sterols and phenols. Bromine as well as iodine seem to be the main single elements. In the antipatharian, during the skeletal formations the dominant structural components are represented by chitin and an antipathin, some kind of halogenated scleroprotein. Antipathin appears to be related with specific material properties of skeletons of black coral. The antipatharians skeletons are less rigid and more elastic in comparison to another biomaterials used as structural components in the nature such as bone, wood, insect cuticle and mollusc shell. Simultaneously, the density of antipathin is lower than bone or shell and higher than wood, but almost similar to cuticle of arthropods.
  • Chapter
    About 63% of the known antipatharian genera occur at mesophotic depths (30–150 m), with the majority extending into the deep sea. Along the continental shelf and offshore sites, antipatharians tend to increase in diversity and abundance with depth, reaching a peak at mesophotic depths due to favorable environmental factors enhancing their settlement and growth and biotic factors associated with lower levels of competition for space. A review of taxonomic and ecological studies for shallow and mesophotic antipatharians is presented for four regionally based case studies, three in the tropics (1) Central Indo-Pacific, plus adjacent sections of the Western Indo-Pacific, (2) Eastern Indo-Pacific (primarily Hawaiʻi), and (3) the Caribbean Sea) and one at temperate latitudes in the Mediterranean Sea and adjacent sections of the Northeast Atlantic. The mesophotic fauna is mainly represented by the families Antipathidae, Aphanipathidae, and Myriopathidae. The most diverse community is found in the Central/Western Indo-Pacific, followed by the Caribbean Sea. The tropical antipatharians are represented by shallow species that extend their distribution into the upper mesophotic zone (30–60 m), while the temperate antipatharians consist of deepwater (> 150 m) species that extend upward into the lower part of the mesophotic zone. Black corals in mesophotic coral ecosystems can be habitat-forming components of benthic assemblages on hard substratum. They have an enormous potential for hidden biodiversity and play an important ecological role for the broader marine ecosystem. The threats to antipatharians consist of demersal fishing activities and coral harvesting, which may be highly destructive to these poorly understood systems.
  • Article
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    There are more coral species in deep, cold-waters than in tropical coral reefs. This broad-ranging treatment is the first to synthesise current understanding of all types of cold-water coral, covering their ecology, biology, palaeontology and geology. Beginning with a history of research in the field, the authors describe the approaches needed to study corals in the deep sea. They consider coral habitats created by stony scleractinian as well as octocoral species. The importance of corals as long-lived geological structures and palaeoclimate archives is discussed, in addition to ways in which they can be conserved. Topic boxes explain unfamiliar concepts, and case studies summarize significant studies, coral habitats or particular conservation measures. Written for professionals and students of marine science, this text is enhanced by an extensive glossary, online resources, and a unique collection of color photographs and illustrations of corals and the habitats they form. © J. Roberts, A. Wheeler, A. Freiwald and S. Cairns 2009 and Cambridge University Press, 2009.
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    Count rates, representing the rate of 14 C decay, are the basic data obtained in a 14 C laboratory. The conversion of this information into an age or geochemical parameters appears a simple matter at first. However, the path between counting and suitable 14 C data reporting (table 1) causes headaches to many. Minor deflections in pathway, depending on personal interpretations, are possible and give end results that are not always useful for inter-laboratory comparisons. This discussion is an attempt to identify some of these problems and to recommend certain procedures by which reporting ambiguities can be avoided.
  • Article
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    Black corals (order Antipatharia) are important long-lived, habitat-forming, sessile, benthic suspension feeders that are found in all oceans and are usually found in water depths greater than 30 m. Deep-water black corals are some of the slowest-growing, longest-lived deep-sea corals known. Previous age dating of a limited number of black coral samples in the Gulf of Mexico focused on extrapolated ages and growth rates based on skeletal Pb-210 dating. Our results greatly expand the age and growth rate data of black corals from the Gulf of Mexico. Radiocarbon analysis of the oldest Leiopathes sp. specimen from the upper De Soto Slope at 300 m water depth indicates that these animals have been growing continuously for at least the last 2 millennia, with growth rates ranging from 8 to 22 mu m yr(-1). Visual growth ring counts based on scanning electron microscopy images were in good agreement with the C-14-derived ages, suggestive of annual ring formation. The presence of bomb-derived C-14 in the outermost samples confirms sinking particulate organic matter as the dominant carbon source and suggests a link between the deep-sea and surface ocean. There was a high degree of reproducibility found between multiple discs cut from the base of each specimen, as well as within duplicate subsamples. Robust C-14-derived chronologies and known surface ocean C-14 reservoir age constraints in the Gulf of Mexico provided reliable calendar ages with future application to the development of proxy records.
  • Article
    The Age Calibration Program, CALIB, published in 1986 and amended in 1987 is here amended anew. The program is available on a floppy disk in this publication. The new calibration data set covers nearly 22 000 Cal yr (approx 18 400 14C yr) and represents a 6 yr timescale calibration effort by several laboratories. The data are described and the program outlined. -K.Clayton
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    A model was developed that can be used to analyze the relationship between body size of indeterminately growing animals and habitat suitability. The model states that the amount of energy available for reproduction in a given season is maximized by maximizing the difference between energy intake and energetic cost as functions of individual mass. Two problems of energy allocation to growth and reproduction are considered as different forms of the model, one where space to store energy during the season is limiting within the organism and a second where it is not. The general model predicts that optimum size will increase with habitat suitability (more prey or lower physiological stress) and that asexual reproduction (colony or clone formation) can often be more energetically favorable than growth to the largest possible individual body size. Relationships between body size, morphology, prey capture, and energetic cost were examined for a passive suspension feeder, the sea anemone Anthopleura xanthogrammica in Washington State. Prey capture was measured in the field for naturally occurring prey and for marked prey released in field experiments. Energetic cost was measured as mass loss during starvation in aquaria. Results show that: (1) prey capture depends on the surface area of the feeding structure, (2) growth isometric in this species, and (3) energetic cost increases as a higher power of mass than does prey capture. Observed patterns of size distribution and prey capture rate in the field are as predicted by the model. Larger individuals are found in habitats having more prey or lower stress. For example, Anthopleura xanthrogrammica increases in size from the high to the low intertidal. Low intertidal habitats have both more feeding time and generally lower summer temperatures and are thus more energetically suitable than those in the high intertidal.
  • Article
    Deep-water (307 697 m) antipatharian (black coral) specimens were collected from the southeastern continental slope of the United States and the north-central Gulf of Mexico. The sclerochronology of the specimens indicates that skeletal growth takes place by formation of concentric coeval layers. We used 210Pb to estimate radial growth rate of two specimens, and to establish that they were several centuries old. Bands were delaminated in KOH and analyzed for carbon and nitrogen stable isotopes. Carbon values ranged from -16.40/00 to -15.70/00; the oldest specimen displayed the largest range in values. Nitrogen values ranged from 7.70/00 to 8.60/00. Two specimens from the same location and depth had similar 15N signatures, indicating good reproducibility between specimens.
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    Complex biogenic habitats formed by corals are important components of the megabenthos of seamounts, but their fragility makes them susceptible to damage by bottom trawling. Here we examine changes to stony corals and associated megabenthic assemblages on seamounts off Tasmania (Australia) with different histories of bottom-contact trawling by analysing 64 504 video frames (25 seamounts) and 704 high-resolution images (7 seamounts). Trawling had a dramatic impact on the seamount benthos: (1) bottom cover of the matrix-forming stony coral Solenosmilia variabilis was reduced by 2 orders of magnitude; (2) loss of coral habitat translated into 3-fold declines in richness, diversity and density of other megabenthos; and (3) megabenthos assemblage structures diverged widely between trawled and untrawled seamounts. On seamounts where trawling had been reduced to <5% a decade ago and ceased completely 5 yr ago, there was no clear signal of recovery of the megabenthos; communities remained impoverished comprising fewer species at reduced densities. Differences in community structure in the trawled (as compared to the untrawled) seamounts were attributed to resistant species that survived initial impacts, others protected in natural refugia and early colonisers. Long-term persistence of trawling impacts on deep-water corals is consistent with their biological traits (e.g. slow growth rates, fragility) that make them particularly vulnerable. Because recovery on seamounts will be slow, the benefits from fishery closures may not be immediately recognisable or measureable. Spatial closures are crucial conservation instruments, but will require long-term commitments and expectations of performance whose time frames match the biological tempo in the deep sea.
  • Article
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    The radial growth rates and ages of three different groups of Hawaiian deep-sea 'corals' were determined using radiocarbon measurements. Specimens of Corallium secundum, Gerardia sp., and Leiopathes glaberrima, were collected from 450 {+-} 40 m at the Makapuu deep-sea coral bed using a submersible (PISCES V). Specimens of Antipathes dichotoma were collected at 50 m off Lahaina, Maui. The primary source of carbon to the calcitic C. secundum skeleton is in situ dissolved inorganic carbon (DIC). Using bomb ¹C time markers we calculate radial growth rates of 170 m y¹ and ages of 68-75 years on specimens as tall as 28 cm of C. secundum. Gerardia sp., A. dichotoma, and L. glaberrima have proteinaceous skeletons and labile particulate organic carbon (POC) is their primary source of architectural carbon. Using ¹C we calculate a radial growth rate of 15 m y¹ and an age of 807 {+-} 30 years for a live collected Gerardia sp., showing that these organisms are extremely long lived. Inner and outer ¹C measurements on four sub-fossil Gerardia spp. samples produce similar growth rate estimates (range 14-45 m y¹) and ages (range 450-2742 years) as observed for the live collected sample. Similarly, with a growth rate of < 10 m y¹ and an age of 2377 years, L. glaberrima at the Makapuu coral bed, is also extremely long lived. In contrast, the shallow-collected A. dichotoma samples yield growth rates ranging from 130 to 1,140 m y¹. These results show that Hawaiian deep-sea corals grow more slowly and are older than previously thought.
  • Article
    The amount of 14C produced by nuclear bomb testing that entered the Atlantic Ocean by late 1972 was 1.71×10-8 mumol/cm2 of ocean surface area for the west Atlantic (36°S-45°N) and 1.18×10-8 mumol/cm2 for the east Atlantic (50°S-28°N) Geochemical Ocean Sections Study stations. There are strong latitudinal differences in the integrated amount of bomb 14C content in Atlantic waters. Bomb-produced 14C is mostly encountered near the center of the large mid-latitude gyres, whereas the equatorial region has a lower 14C inventory. The average ocean wide vertical distribution of bomb 14C in the Atlantic can be explained by a vertical eddy diffusion coefficient of 4.0 cm2/s in the surface mixed layer plus thermocline gyre reservoirs. The average 14C activity per unit area measured in the Atlantic yields an atmosphere-ocean CO2 exchange rate of 23 mol/m2 yr, which is equivalent with an atmospheric CO2 residence time of 6.8 years.