P.S.Z.N.

: Marine Ecology, 18 (I): 83-94 (1997)

0 1997 Blackwell Wissenschafts-Verlag, Berlin Accepted: October 4, 1996
ISSN 0173-9565

Life in the Slow Lane: Growth and

Longevity of Cold-seep
Vestimentiferans
C. R.FISHER,I. A. URCWO,M. A. SIMPKINS*
& E. NIxt

The Pennsylvania State University, Department of Biology, 208 Mueller Laboratory,

University Park, PA 16802, USA.

With 2 figures and 2 tables

Key words: Vestimentiferan, cold seep, growth, age, deep sea.

Abstract. Yearly, in siru growth of two species of cold-seep vestimentiferan tubeworms was measured
using a combination of banding and video analysis. A total of 53 animals from 15 different aggregations
were included in the study, which was conducted over a Cyear period. The most common species,
Lamellibrachiu sp., grows very slowly (averaging 0.77 cm .a-') and yet commonly reaches lengths over
2 m. Based on conservative calculations we conclude that individuals in mature aggregations are a
minimum of 100 a old and are likely to be much older. Smaller numbers of a second species of
vestimentiferan (related to escarpids) were also monitored over this period. We consider it likely that
the individuals of the second species banded for this study were already on the plateau of their growth
curve, and that the very low average growth rates recorded here would lead to an overestimate of the
ages of the individuals used in this study. However, life history considerations combined with the
extremely low measured growth rates of this species suggest it is also long-lived.

Problem
Vestimentiferan tubeworms are one of the dominant faunal types at many hydro-
thermal vents and cold seeps in the deep sea (HESSLER& SMITHEY, 1983;
TUNNICLIFFE, 1991; LUTZ & KENNISH,1993). When these communities were dis-
covered less than 2 decades ago they were largely considered as interesting, but
isolated, oddities. However, the widespread discoveries of numerous additional
communities in the last 15 years suggest that these communities are pandemic in
the deep sea and raise questions concerning the magnitude and impact of vent and
seep primary production to the nutrient-limited ambient deep-sea fauna (GAGE&
TYLER,1991). As a result of their symbiosis with chemoautotrophic sulfide-oxi-
dizing bacteria, vestimentiferans can be considered one of the primary producers
in the vent and seep communities (FISHER, 1990; CHILDRBS& FISHER,1992;
GARJXNER& JONES,1993). Furthermore, as a result of their size and abundance,

Currently at the University of Alaska, Fairbanks, USA.

t Currently at South Carolina Coastal University, USA.

U. S.Copyright Clearance Center Code Statement: 0 173-9565/97/ 1801-0083$14.00/0

84 FISHER,
URCUYO, SIMPKINS
& NIX

they provide a habitat for numerous associated species of smaller animals (MAC-
DONALD et af., 1989; CARNEY, 1994). In short, vestimentiferans are key ecosystem
structuring species in abundant, widespread and productive deep-sea hydrothermal
vent and cold-seep environments.
Vestimentiferan tubeworms are also one of the pioneer metazoan species to
colonize a new hydrothermal vent site, and can quickly grow to dominate their
habitats (FUSTECet af., 1987; HESSLER et af., 1988; LUTZet af., 1994). Very high
growth rates have been measured or inferred for a variety of hydrothermal vent
vestimentiferans (FUSTECet af., 1988; Roux et af., 1989; TUNNICLIFFE et af., 1990;
LUTZet al., 1994). Recently, a study based on video analysis of the establishment
and growth of a hydrothermal vent community on the East Pacific Rise documented
growth of Riftiapachyptifa tubes to lengths greater than 1.5m (and sexual maturity)
in less than 2 years ( L m z et af., 1994). The authors suggest that this represents the
fastest rate of linear growth documented for any marine invertebrate. Although
this suggestion is reasonable, it should be noted that tube growth does not
reflect tissue growth in this species because the animal does not inhabit its entire
tube.
Due to the ephemeral nature of hydrothermal vents, a life history which involves
fast growth to reproductive size has obvious advantages to fauna strictly dependent
on the vent effluent for nourishment (FUSTECet af., 1987; TUNNICLIFFE, 1991; LUTZ
et al., 1994). However, the cold-seep environment on the Louisiana Slope in the
Gulf of Mexico is much more stable than the hydrothermal vent environment. Salt
and hydrocarbon seepage in the Gulf of Mexico is driven largely by salt tectonics
(SALVADOR, 1991). Here, the sea floor is covered by an average depth of over 1000
m of sediment which buffers and augments an already slow and relatively stable
process (COLEMAN et af., 1991; SALVADOR, 1991). In addition to sulfide advecting
from deep thermogenic and biogenic sources, sulfide is also produced biogenically
in surficial sediments (CARNEY, 1994). Thus, unlike many hydrothermal vents where
individual vents persist for years to decades (FUSTECet al., 1987; HESSLER et af.,
1988; LUTZ& KENNISH, 1993), the Gulf of Mexico seeps may provide an environ-
ment rich in sulfide for centuries.
Two anecdotal observations had suggested that vestimentiferans on the Lou-
isiana Slope of the Gulf of Mexico might be long lived. One was the collection of
seep vestimentiferans with large encrustations of deep-sea sponges and octocorals
on the tubes near their anterior (growing) end, suggesting that they had lived a
long time after reaching moderately large sizes (FISHERet af., 1990). The other was
the collection of many seep vestimentiferans which had the attached posterior ends
of the tubes buried by over a meter of sediment (pers. obs.). Based on what is
known of the life history of these animals, the motile larvae attach to a solid
substrate and the animals are sessile for the rest of their life (SOUTHWARD, 1988;
JONES& GARDINER, 1989; GARDINER & JONES, 1993). Therefore the sediment over
the rock on which they originally settled must have accumulated over the life span
of the individuals. Consistent with this thesis are observations of newly settled seep
vestimentiferans on exposed rocks, and never in sediments. In this area of the Gulf
of Mexico, the background sedimentation rate is on the order of 1-6 cm per
hundred years and although sedimentation is undoubtedly enhanced by the bush-
like structure of the tube-worm aggregations, burial by 1 m of sediment implies at
least moderately old animals (MACDONALD et af., 1989). Here we report on direct,
Growth of cold-seep vestimentiferans 85
in situ growth measurements of two species of cold-seep vestimentiferans over four,
1-year periods.

Material and Methods

1. The vestimentiferans

Two species of vestimentiferan are found in association with hydrocarbon and saline seeps at depths
less than 1000 m on the Louisiana Slope of the Gulf of Mexico. Neither has been adequately described
to date. However, the presence of two species is widely recognized in over 10 years of ecological and
physiological literature because they are easy to differentiate using external characters. One species was
immediately recognized as belonging in the genus Lamellibrachia (JONES,1985) and must still be referred
to as Lumellibrachia sp. until a detailed comparison of the lamellibrachid species can be completed (S.
GARDINER, Bryn Mawr College, USA,pers. comm.). The second species belongs in a new genus and
perhaps a new family related to the family Escarpidue (S. GARDINER, Bryn Mawr College, USA, pers.
comm.). In previous literature this species has been variously referred to as ‘escarpid-like’ or ‘the
escarpid’. The two species usually occur in mixed aggregations dominated by the longer lamellibrachids,
with the escarpid-like species found around the base of the aggregations (Fig. la). In contrast to
aggregations of large individuals, numerous individuals of the escarpid-like species are often present in
aggregations of small, presumably young, vestimentiferans (Fig. 1b). Due to physical limitations on
banding very small animals with the device employed, we selectively banded medium- to large-sized
animals for this study. The 53 individuals monitored for this study come from 15 different aggregations
distributed at three different sites between 540 and 730 m depth. The three sites (Bush Hill, G C 234,
and G C 272) are described in NIXet a/. (1995) and MACDONALD et al. (1989, 1990).

2. In situ banding

To determine growth rates for these species, a device was constructed which deployed plastic cable ties
around the tubes near their anterior end. The hydraulically actuated device deployed up to eight bands
per dive to a pre-set tension around the animals, and released the bands after deployment. Attached to
each band were coloured and numbered tags for identification of the individual vestimentiferans year
after year. The device was positioned by the pilot of a Johnson Sea-Link submersible such that the loop
of a band surrounded the anterior end of a vestimentiferan tube, and then actuated to secure the band
around the tube. The position of the band with respect to the growing end of the worm was then
documented by video. Over the course of this study, 37 lamellibrachids and 16 individuals of the
escarpid-like species were banded with adequate video documentation for inclusion in this study.

3. Determination of yearly growth increment

Three methods were used to determine the yearly growth increment of the seep vestimentiferans.
Whenever possible, direct measurements were made from digitized video images in 2 or more years.
For these measurements, the outer surface of the band itself was used to provide scale in the images
and allow determination of the distance between the band and the anterior end of the tube. Since the
band lies flat on the tube’s surface and is approximately perpendicular to the tube’s long axis, measure-
ment of the band width compensates for the angle of documentation (Fig. 2). Re-video in subsequent
years was used to determine the yearly in situ growth increments of the tubes of individual animals.
In cases where direct videoscopic measurements were not possible due to poor image quality, tube
elongation rates were estimated based on the number of growth rings deposited and the average distance
between rings. The average distance between growth rings for each species was determined by measuring
86 FISHER,
URCWO, SIMPKINS & NIX

Fig. 1. (a) A typical mature aggregation of vestimentiferan tubeworms at a shallow seep site in the Gulf
of Mexico. The clearly visible vestimentiferans are all lamellibrachids. and only a few escarpids are
barely discernible around the base of the aggregation (arrows). Approximately 1 to 1.5 m of the tubes
of the worms are visible above the sediment in this aggregation. (b) A small aggregation of small
worms and mussels. Although the lamellibrachids dominate this young aggregation, many escarpid-
like vestimentiferans are easily recognized by the flared ends of their tubes.
Growth of cold-seep vestimentiferans 87

Fig. 2. Digitized video image of a banded lamellibrachid tube worm. Measurements made offa digitized
image are represented by Xi (pixels in the new terminal increment(s)) and Xt (pixels in the band). These
are converted to actual Li (length of terminal increment in mm) based on Lt (known width of the band).

the distance between each of the anterior 10-15 rings on the tubes of 20 lamellibrachids and 14 of the
escarpid-like species (a total of 239 and 172 rings were measured, respectively) (SIMPKINS,
1994).
Fourteen banded lamellibrachids and seven banded escarpid-like vestimentiferans were collected to
verify the efficacy of the method and determine the approximate size range of the banded vesti-
mentiferans. Thirteen of these animals were adequately documented by video in previous years (for
measurements or ring counts) and the growth increment for those individuals was determined by direct
measurement of the distance between the band and the end of tubes after collection.

4. Morphometrics

The lengths of the tubes of the collected vestimentiferans were determined from the anterior end to a
point on the posterior end where the exterior tube diameter had decreased to 2 mm. This was done by
laying a string along the irregularly shaped tube and then measuring the length of the string. Portions
of the tube posterior to this point were often not recovered because during collections the posterior end
of the tube normally breaks off near its point of attachment to buried carbonate rocks. We calculate
that the tissue contained in the portion of the tube less than 2 mm in diameter represents at most 3%
of the biomass of any animal collected for this study. This calculation is based on the measured biomass
of the smallest banded animal collected (to the point where the exterior tube diameter decreased to 2
mm), and the generous assumption that the portion less than 2 mm in diameter was 30 cm long and
filled completely with tissue.
88 FISHER,
URCUYO, SIMPKINS
& NIX

Results
The data on tube growth of each individual over the 4 years of this study are shown
in Tables 1 and 2. The differences in the amount of data collected from different

Table I . Growth of Lumellibrachia sp. in the Gulf of Mexico between 1991 and 1995.

growth (rings, mm)

91-92 92-93 93-94 9495

11 10 15 11 average
site group banded collected months months months months [mm * a-’1
-

Bush Hill BH 1 1991 1994 I. 7 - - 7

BH I 1991 1993 5, 25 4.28 26
BH2 1991 1993 3, 16 0 8
BH3 1991 1993 0 0 0
BH5 I992 NC - - 1,5 5.6
BH6 1992 NC - 4,26 - 20
BH6 1992 NC - 0 - 0
BH7 1993 NC - I, 7 7.6
BH7 1993 NC 4,22 - 17
BH7 1993 NC 2,9 - 7
BH7 1993 NC 3,20 2, 1 1 13.8
BH7 1993 NC 6, 22 6, 29 24.7
BH7 1993 NC 2, I4 3,2I 18.2
BH8 1993 NC I,7 1,7 6.9
BH8 1993 NC 0 0 0
BH8 1993 NC 0 - 0
BH8 I993 NC 0 0 0
GC 234 GC234- 1 1991 NC 3, 18 2, 17 5.35 I. 7 17.7
GC234-1 1991 NC 0 0 - - 0
GC234-1 1991 NC 0 0 0 - 0
GC234-1 1991 NC 0 0 2,8 2, 12 4.9
GC234-2 1991 1992 1,3 3
GC234-2 1991 1992 3, 21 20
GC234-2 1991 1992 0 0
GC234-2 1991 I992 0 0
GC234-3 1993 NC 1,8 - 6.5
GC234-3 1993 NC 0 - 0
GC234-4 1993 NC - 0 0
GC234-4 1993 NC 0 0 0
GC234-4 1993 NC 0 2, 15 8.3
GC234-5 1993 NC 1,9 I. 7 7.3
GC234-5 1993 NC 3, 14 - 11
GC234-6 1993 NC 173 1,4 3.3
GC234-6 1993 NC I, 9 I. 7 7.4
GC234-6 1993 NC 3,24 2, 1s 17.3
02234-6 1993 NC 2, I4 3, 20 17.8
GC234-6 1993 NC 0 - 0

NC = not collected as of 1995.

Italics denote growth that was calculated using ring counts and the average ring length of the species.
A dash (-) shows years of no growth measurement due to the lack of adequate video for image analysis.
Growth of cold-seep vestimentiferans 89
Table 2. Growth of the escarpid-like vestimentiferan in the Gulf of Mexico between 1991 and 1995.

growth (rings, mm)

91-92 92-93 93-94 94-95

11 10 15 11 average
site group banded collected months months months months [mm. a-'1

Bush Hill BH2 1991 1993 0 0 0

BH2 1991 1993 0 0 0
BH2 1991 1993 0 0 0
BH2 1991 1993 0 0 0
BH3 1991 1993 I , I1 0 5.5
BH4 1992 NC 0 I , I1 6.5
BH4 1992 NC 0 0 0
BH5 1992 NC - - 0 0
BH5 1992 NC 0 1, I I 0 2.8
BH5 1992 NC 0 0 0 0
BH5 1992 NC 0 0 0 0
GC 234 GC234-3 1993 NC 0 - 0
GC 272 GC234-8 1993 NC 0 - 0
GC234-8 1993 NC - 0 0
GC234-8 1993 NC - 0 0
GC234-8 1993 NC 0 0 0

NC = not collected as of 1995.

Italics denote growth that was calculated using ring counts and the average ring length of the species.
A dash (-) shows years of no growth measurement due to the lack of adequate video for image analysis.

individuals are a result of variation in the duration of the experiments and in the
quality and quantity of the video documentation of the banded animals in each
year. Anterior tube growth was documented in 24 of the 37 lamellibrachids over
at least one of the 4 years. Growth was quite variable both between individuals
and, in some cases, between years by an individual. Six individuals did not grow
for 2 years in a row and another did not grow over 3 consecutive years. The
highest yearly average for a lamellibrachid was 26 mm. The average yearly growth
increment among all lamellibrachids was 7.7 mm (SD = 8.0, n = 37) and the
median growth increment was 6.9 mm. The 14 banded lamellibrachids which were
collected ranged in tube length from 98 to 216 cm.
Of the individuals of the escarpid-like species monitored, only three grew in any
1 year and all passed at least 1 year without growing. Their highest yearly average
for an individual was 6.5 mm, their species average yearly growth increment was
0.9 mm (SD = 2.1, n = 16). However, this average is heavily influenced by the large
number of individuals which did not grow over the course of this study, and in
fact the median yearly growth increment for this species is 0 mm. The seven banded
escarpid-like individuals which were collected ranged in tube length from 47 to 104
cm.
The average ring spacing on the tubes for the two species was significantly
different (P < 0.0001, 2-sample t-test). The average distance between rings on
the lamellibrachid tubes was 6.7 mm (SD = 3.0, n = 239 measurements from 20
90 FISHER,
URCWO,SIMPKINS
& NIX

individuals). The average distance between rings on the tubes of the escarpid-like
vestimentiferan was 11.4 mm (SD= 4.3, n = 172 measurements from 14 indi-
viduals). There were significant differences in average ring spacing between some
individuals of the same species, so, for determination of the relation of the terminal
increment to previous ring spacing, comparisons were made within an individual.
The terminal increment in all 14 of the escarpid-like tubes measured was within
the 95 % confidence interval of the mean of the adjacent 10 increments, suggesting
that the terminal increment was not in the process of extension. Similar analysis
indicated that the terminal increment of 17 of the 20 lamellibrachids had reached
its full length.

Discussion
The seep vestimentiferans are anatomically and physiologically quite similar to
their hydrothermal vent relatives (FISHERet al., 1988; FISHER,1990). However,
unlike the most massive vent species, Rijtiapachyptila, living tissue extends virtually
the full length of the tube, tapering to a hair-width near the point of original tube
attachment (GARDINER & JONES, 1993); thus, tube growth is a direct reflection of
living-tissue growth in these species. There are suggestions that some ves-
timentiferan species are capable of extending their tube from the posterior end.
Our observations of tangled, small-diameter, transparent tubes posterior to a point
of attachment to the substrate in the seep species suggest that this may be true for
them as well. However, because of the very small diameter of the animal at the
posterior end, even very significant amounts of linear extension from the posterior
end would result in a very minimal increase in biomass in these species. Further-
more, because we did not measure tube length beyond the point that it reaches 2
mm in external diameter, and because we did not consider the first 50 cm of linear
growth (up to a tube diameter of about 5 mm) in our age estimates, the possibility
of posterior tube elongation does not affect our age estimates.
Over the first 2 years of this study about half of the Lamellibrachia sp. monitored
elongated their tube (and the other half did not). Between 1994 and 1995, the
proportion elongating their tube increased to 78%. One individual in this study
elongated its tube every year for 4 years at an average rate of 18 m m - a - ' . Another
individual from the same aggregation did not elongate over any of the 3-year
periods it was observed. Other animals in that aggregation and elsewhere grew in
some years, but not in others (Table 1). This kind of variability in growth within
an aggregation was typical, although aggregations were monitored in which either
all, or none, of the tagged individuals grew in a given year. Thus, tube elongation
is episodic in this species and variation in growth rate is patchy on several spatial
scales as well.
Based on the full data set of 37 lamellibrachids from 13 different aggregations
monitored over 4 years (Table l), it is apparent that the lamellibrachids grow at a
much slower rate than their hydrothermal vent relatives. Even the largest yearly
growth increment recorded is over an order of magnitude less than hydrothermal
vent vestimentiferans. However, these seep species can reach body lengths more
than twice that of the largest documented Rijtia pachyptila (the largest known
Growth of cold-seep vestimentiferans 91
hydrothermal vent species) (JONES, 1981). Individuals of Lamellibrachia sp. in
excess of 3 m in length have been collected several times in the past even though
efforts are made to avoid collecting larger individuals because they d o not fit in the
submersible collection box (pers. obs.).
The total lengths of most of the banded worms used in this study are unknown
because they are still in situ, partially buried with sediment. The banded lamel-
librachids collected to date ranged from 98-216 cm in length. The collected indi-
viduals were banded in 1991. During the 1993 field season we selectively banded
smaller individuals. Thus, it is reasonable to expect that our data covers an even
wider size range of animals (none of the animals banded in 1993 have been collected
and we are still monitoring their growth). Within the size range of banded animals
there is no indication of a decrease in growth rate with increasing size. In fact, the
largest banded individual collected (2 16 cm) added 20 mm of tube length (3 growth
rings) between 1991 and 1992, and was thus among the fastest growing animals in
this study. However, due to physical limitations in banding worms with very small
diameter tubes, we have no data on lamellibrachids with anterior tube diameter
less than about 5 mm (and tube lengths less than approximately 50 cm). It is very
likely that the smaller individuals would have a significantly higher growth rate,
due to the allometry of metabolism and growth. Even if the time for growth up to
the moderate sizes of the banded individuals monitored in this study is assumed to
be instantaneous, the very slow subsequent growth rates, and the total lengths
attained by these lamellibrachids, indicate that they are a long-lived species. A
-
simple calculation, assuming an average tube elongation rate of 1 cm a-' (greater
than measured in this study), suggests that it takes a lamellibrachid 150 years to
grow from 50 to 200 cm in tube length. Even calculations based on the highest
growth rates measured indicate that the mature vestimentiferan aggregations which
abound in this area are at least 100 years old.
Only three of the 16 escarpid-like vestimentiferans monitored elongated their
tubes in any year (Table 2). Over the course of this study it was often observed
that although this species is abundant in the relatively rare aggregations of very
small/young individuals ( < 30-50 cm), they are sparse and only found at the basal
periphery of the much more common larger aggregations of moderate- to large-
sized worms (Fig. 1). This observation suggests that the escarpid-like species may
have a life history very different from Lamellibrachia sp., and that the former may
be a pioneer species in the seep vestimentiferan habitat. Thus, we consider it likely
that we have preferentially sampled the late stages of the escarpid-like species life
cycle. Further studies are currently underway to address this issue and it is prema-
ture to make estimates of the ages and life spans of this species from the banding
data. However, recruitment of all individuals (both species) present within a par-
ticular aggregation is likely to occur over a relatively limited time span and end
when the recruitment substrate becomes buried by sediment (SIMPKINS, 1994).
This suggests that both species present in the large aggregations dominated by
Lamellibrachia sp. are of comparable age. The very low average growth rates and
shorter tubes documented for the escarpid-like species are consistent with this
hypothesis.
Both species of seep vestimentiferan have visible external rings on their tubes
(Figs 1 and 2), and it was hoped that these rings might assist in determining the
ages of individual seep vestimentiferans. However, the variability in the number of
92 FISHER,
URCWO, SIMPKINS
& NIX

rings deposited by different individuals in a single year (Table 1) indicates that

there is no simple temporal meaning to the rings, and they are therefore of little
use for aging animals. Based on examination of collected specimens and analysis
of video footage, we agree with GARDINER & JONES (1993) that the rings mark the
increment of tube deposited in a single growth ‘spurt’, and that rings on the sides
of the tubes represent the relic of the flared tube ends. The flared ends strengthen
the terminus of the tube and allow the worm to extend and retract without catching
on the end of the tube. The terminal tube increment of many of the individuals
collected was noticeably thinner walled than adjoining proximal sections, indicating
that the second step in the deposition of a new tube increment is thickening the
wall. We suggest that the entire increment is deposited over a relatively short
period, and then strengthened over an additional, longer period. This is supported
by the fact that no increases in the original terminal increments were detected on
any of the banded animals measured in situ and that the terminal increment of 31
of 34 tubes analysed was within the 95% confidence interval of the mean of the
adjacent 10 increments.
The longevity and low growth rates of the cold-seep vestimentiferans reflect life
history strategies very different from their hydrothermal vent relatives. The deep
sea in general is a relatively stable environment (GAGE& TYLER,1991). In the
case of the seep vestimentiferans, the energy source for their chemoautotrophic
symbionts is stable and constant over long periods. However, sulfide is not as
abundant around the seep vestimentiferans as it is in hydrothermal vent ves-
timentiferan habitats. Slow growth and long life are therefore well suited to seep
vestimentiferans. At the other extreme are the hydrothermal vent species: their
fast growth to reproductive size reflects their adaptation to an energy-rich, but
ephemeral, habitat. Thus, within the small phylum Vestimentifera, there are species
which are extremely fast-growing and others which are very long lived.

Summary
Vestimentiferan tubeworms around hydrocarbon seeps in the Gulf of Mexico grow
much more slowly and live much longer than related hydrothermal vent species.
Average. growth rates are under 1 cm per year for Lamellibrachia sp. and these
seep vestimentiferans reach lengths over 2 m. Conservative estimates of the ages
of individuals in mature aggregations range from 100 to 150 years, but they
may live considerably longer. Thus, the life history strategy of these cold-seep
vestimentiferans is very different from that of hydrothermal vent species.

Acknowledgements
The authors would like to acknowledge the following: The captains, pilots and crew of the RV Seward
Johnson, RV Edwin Link and DSRV Johnson Sea Link I and 11, without whose talents the submersible
manipulations necessary for this study would not have been possible; CRISTIETZand Harbor Branch
OceanographicInstitution Engineering division for design and construction of the banding device; IAN
MACDONALD, JENNY VODENICHAR, JILL ZANDE,KEVINKELLY, MECSTREAMS, STEVE GARDINER, NEIL
TAUFEN, MATTSWIDER, KT SCOTTand STEWMACKENZIE for assistance at sea and in the laboratory.
This research was supported by The Mineral Management Service, The National Science Foundation,
Growth of cold-seep vestimentiferans 93
The National Oceanic and Atmospheric Administration's National Undersea Research Program and
the Harbor Branch Oceanographic Institution.

References

CARNEY, R. S., 1994: Consideration of the oasis analogy for chemosynthetic communities at Gulf of
Mexico hydrocarbon vents. Geo-Mar. Lett., 1 4 149-159.
CHiLDRm, J. J. & C. R. FISHER, 1992: The biology of hydrothermal vent animals: physiology, bio-
chemistry and autotrophic symbiosis. Oceanogr. Mar. Biol. Annu. Rev., 30:3 3 7 4 1 .
COLEMAN, J. M., H. H. ROBERTS& W. R. BRYANT,1991: Late quarternary sedimentation. In: A.
SALVADOR (Ed.), The Gulf of Mexico Basin. The Geology Society of America, Boulder, CO: 3 2 5
352.
FISHER,C. R., 1990: Chemoautotrophic and methanotrophic symbioses in marine invertebrates. Rev.
Aquat. Sci., 2 399436.
--, J. J. CHILDRESS & N. K. SANDERS, 1988: The role of vestimentiferan hemoglobin in providing an
environment suitable for chemoautotrophic sulfide-oxidizing endosymbionts. Symbiosis, 5: 229-

--. 246.
M. C. KENNICUTT I1 & J. M. BROOKS, 1990: Stable Carbon isotopic evidence for carbon limitation
in hydrothermal vent vestimentiferans. Science, 247: 1094-1096.
FUSTEC,A,, D. DESBRUYERES & S. KIM JUNIPER, 1987: Deep-sea Hydrothermal Vent communities at
13"Non the East Pacific Rise: Microdistribution and temporal variations. Biol. Oceanogr., 4: 121-
164.
--, D. DESBRUYERES & L. LAUBIER, 1988: Biomass estimation of animal communities associated with
deep-sea hydrothermal vents near I3"N/EPR. Oceanologica Acta, spec. vol., 8 15-22,
GAGE,J. D. & P. A. TYLER,1991: Deep-sea Biology: A Natural History of Organisms at the Deep-sea
Floor. University Press, Cambridge; 504 pp.
GARDINER, s. L. & M. L. JONES,1993: Vestimentifera. In: s. L.GARDINER (Ed.), Microscopic Anatomy
of Invertebrates. Volume 12: Onychophora, Chilopoda, and Lesser Protostomata. Wiley-Liss,
New York: 371460.
HESSLER, R. R. & W. M. SMITHEY, Jr., 1983: The distribution and community structure of megafauna
at the Galapagos Rift hydrothermal vents. In: P. A. RONA,K. BOSTR~M, L. LAUBIER & K. L.
SMITH,Jr. (Ed.), Hydrothermal processes at seafloor spreading centers. Plenum Press, New York:
735-770.
--, W. M. SMITHEY, M. A. BOUDRIAS, C. H. KELLER, R. A. LUTZ& J. J. CHILDRESS, 1988: Temporal
change in megafauna at the Rose Garden hydrothermal vent. Deep-sea Res., 35: 1681-1710.
JONES, M. L., 1981: Rrfi/iupuchyp/ilu,new genus, new species, the vestimentiferan tubeworm from the
Galapagos Rift geothermal vents. Proc. Biol. SOC.Wash., 93: 1295-1313.
--, 1985:On the Vestimentifera, new phylum: six new species, and other taxa, from hydrothermal vents
and elsewhere. Bull. Biol. SOC.Wash., 6 117-158.
-- & S. L. GARDINER. 1989: On the early development of the vestimentiferan tube worm Ridgeio sp.
and observations on the nervous system and trophosome of Ridgeiu sp. and Rif/iapuc/iyp/i/u.Biol.
Bull., 177: 254-276.
LUTZ,R. A. & M. J. KENNISH,1993: Ecology of deep-sea hydrothermal vent communities: A review.
Rev. Geophys., 31: 21 1-242.
--, T. M. SHANK, D. J. FORNARI. R. M. HAYMON, M. D. LILLEY, K. L. VONDAMN & D. DESBRUYERES.
1994: Rapid growth at deep-sea vents. Nature, 371: 663-664.
MACDONALD, I. R.,G. s. BOLAND, J. s. BAKER, J. M. BROOKS, M. c. KENNICUTT 11 & R. R. BIDIGARE,
1989: Gulf of Mexico hydrocarbon seep communities. 11. Spatial distribution of seep organisms
and hydrocarbons at Bush Hill. Mar. Biol., 101: 235-247.
--. N. L. GUINASSO, J. F. REILLY, J. M. BROOKS,W. R. CALLENDER & S. G. GABRIELLE, 1990: Gulf of
Mexico hydrocarbon seep communities: VI. Patterns in community structure and habitat. Geo-
Mar. Lett., 1 0 244-252.
NIX, E. R., C. R. FISHER,J. VODENICHAR & K. M. SCOTT,1995: Physiological ecology of a mussel with
methanotrophic endosymbionts at three hydrocarbon seep sites in the Gulf of Mexico. Mar. Biol.,
122: 605-617.
Roux, M.. M. RIO,E. SCHEIN. R. LUTZ.L. FRITZ & L. RAGONE, 1989: Mesures in siru de la croissance
des bivalves et des vestimentiferes et de la corrosion des coquilles au site hydrothermal de 13"N
(dorsale du Pacifique oriental). C.R. Acad. Sci. Paris, Ser. 111. 308 121-127.
SALVADOR, A., 1991: Introduction. In: A. SALVADOR (Ed.), The Gulf of Mexico Basin. Geological
Society of America, Boulder, Co: 1-12.
94 URCUYO,
FISHER, & NIX
SIMPKINS
SIMPKINS, M. A., 1994: Hydrocarbon seep vestimentiferans: growth and condition in a variable environ-
ment. MS Thesis, Pennsylvania State University; 60 pp.
SOUTHWARD, E. C., 1988: Development of the gut and segmentation of newly settled stages of Ridgeia
(Vestimentifera): Implications for relationship between Vestimentifera and Pogonophora. J. Mar.
Biol. Assoc. U. K., 68: 465487.
TUNNICLIFFE, V., 1991: The biology of hydrothermal vents: Ecology and evolution. Oceanogr. Mar.
Biol. Annu. Rev., 2 9 319-407.
--, J. F. GARRETT & H. P. JOHNSON, 1990: Physical and biological factors affecting the behaviour and
mortality of hydrothermal vent tubeworms (vestimentiferans). Deep-sea Res., 37: 103-125.