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Vent Larval Life Histories
Susan Mills and Lauren Mullineaux
9 September 2008
Introduction
These larval attributes and species life histories are intended to inform LADDER
dispersal models. We selected vent species based on prevalence and importance in
ecosystem, abundance in our samples, and availability of information.
Some of the measurements are from deep-sea species under deep-sea conditions, but
most were obtained from related shallow-water organisms and at temperatures of 1020°C. We expect development to be slower in the deep sea than shallow because the
reduced (typically ~2°C) temperatures are known to slow metabolic rates and
development. For feeding larvae, we also expect slower development due to low food
availability. Swimming speeds of deep-sea larvae may be lower than shallow relatives,
due to lower metabolic rates and increased viscosity of cold water respectively (e.g.,
Young 1995); sinking speeds also may be reduced by the viscosity. Thus the estimates
from shallow species of stage durations and lifespans are likely to be underestimates,
and movement rates are probably overestimates, for deep species.
Archaeogastropods
Snails and limpets (Bathymargarites symplector, Lepetodrilus spp., Ctenopelta porifera,
Cyathermia naticoides) are numerous and well-preserved in our larval samples and
identifiable to the species level. They are known to be important in vent communities,
and show large abundance changes pre- and post-eruption.
Size: Bathymargarites symplector – 0.32 – 0.35 mm
Lepetodrilus spp. – 0.17 – 0.18 mm
Ctenopelta porifera – 0.29 – 0.31 mm
Cyathermia naticoides – 0.23 – 0.24 mm
Sinking Velocity: estimated 1.0 to 1.5 mm/s, based on measurements of similarly sized
opisthobranch lecithotrophic gastropod Alderia at 25°C (Krug 2007)
Swim Speed: estimated 1.2 mm/s, based on Alderia at 25°C (Krug 2007)
Feeding stage: No feeding stage in the plankton, but Warén suggests (via email), many
gastropods may retain swimming ability for some time after they start benthic feeding.
Precompetent stage length: Unknown, but possibly competent on release.
Competent stage length: Unknown, but likely limited by lipid stores because larva is
non-feeding (Pechenik 1987).
Larval lifespan: In archaeogastropods spawning is internal and larvae typically are
lecithotrophic and metamorphose quickly after spawning (e.g., 3 to 4 days at 27.5 C for
Citarium pica; Bell 1992). Some of the vent species may be brooded for part or all of their
larval development (e.g., Bathymargarites in Warén & Bouchet, 1993), or lay
encapsulated eggs. Eulepetopsis vitrea is reported as a broadcast spawner (Tyler et al.
2008). Number of offspring/brood runs from 10’s to 100’s (Tyler et al. 2008). Larval
development times for shallow-water, planktonic, non-feeding gastropod larvae range
from 4 to 21 days (Grantham et al. 2003). Vent species lifespan is probably less than one
month, but they may retain some mobility after settling into benthos
Other attributes: Some peltospirids, including Ctenopelta, have reduced digestive tracts
as adults.
Bythograeid Crabs
Vent crabs, including the abundant B. thermydron and the more rare B. microps. We
collect these larvae in traps, but rarely in pumps. There are multiple anecdotal reports
of megalopae recruiting ‘en mass’, because they are visible by eye. Juvenile
bythograeids were notably abundant after the eruption.
Size: Bythograea thermydron – 1.5 mm zoea, 8 mm megalopa
Bythograea microps – 3 mm megalopa
Sinking Velocity: estimated as 6 to 20 mm/s for similarly sized shallow species (Chia et
al. 1984)
Swim Speed: measured 40 mm/s for megalopa at 2°C (Epifanio et al. 1999); estimated 817 mm/s for zoea (Mileikovsky 1973) or 9.5 to 15 mm/s for zoea and 42 mm/s for
megalopa upwards (Chia et al. 1984)
Precompetent stage length: Larvae brooded until zoea stage (we find brooding adults in
field, also reported in Perovich et al. 2003)
Feeding stage: Yes, megalopa larva and possibly multiple zoeal stages
Competent stage length: Possibly months. B. thermydron megalopae survived for 191
days at room temperature and sea level pressure (Epifanio et al. 1999).
Larval lifespan: Observed as months
Other attributes: B. thermydron zoea have eye pigments that are receptive to blue light
(Jinks et al. 2002) suggesting they may rise into photic zone. Later stages don’t have the
pigments. This species also appears to move out to the periphery of vent sites to release
its brooded zoea (Perovich et al. 2003).
Bathymodiolid Mussel
The vent mussel Bathymodiolus thermophilus is abundant at EPR vents,
and often displaces Riftia tubeworms as communities progress
through successional stages. A congener B. azoricus is found on the
Mid Atlantic Ridge, where it has been cultured. Mussels are
typically broadcast spawners, the trochophore quickly develops a
shell (D-stage), and then adds more shell (P-II stage) as it feeds in
plankton.
Size: ~ 0.10 mm for P-I and 0.40 to 0.45 mm for P-II
Sinking Velocity: Increases throughout development. Chia et al., 1984 report 1.7 mm/s
(D-stage) and 8.3 mm/s (eyed veliger) for Crassostrea
Swim Speed: Mercenaria mercenaria D-stage swim 0.83 to 1.6 mm/s (Mileikovsky 1973);
Crassostrea swim 0. 8 mm/s upward at D-stage, 2.3 mm/s upward at eyed veliger,
horizontal speeds of 0. 3 and 0.8 mm/s for D-stage and eyed veliger respectively (Chia
et al., 1984)
Precompetent stage length: Mytilus edulis trochophore sets initial shell by 20-24 h at 15°
C, PI stage lasts 1-2 d, precompetent (feeding) PII stage lasts 24 – 27 d (Bayne 1964); at
11° C, precompetent stage lasts 38 d (20 d at 16° C) and competent stage is 46 d (25 d at
16° C) (Bayne 1965). With lots of food, the length of competent stage can be extended to
45 d at 16° C (Pechenik et al. 1990). Other studies show similar delay of development at
low temperature and food levels: M edulis develops to D-stage by 32, 42, 104 h at (18, 12
and 6° C), develops to competent stage by 20-80 d, depending on temperature and food
supply (Sprung 1984). Increasing food supply can accelerate development by 2 to 5fold.
Feeding stage: B. thermophilus feeds in PII stage as it accretes shell. M. edulis feeds in
precompetent and part of competent PII stage (Bayne 1965)
Competent stage length: Mussels caught in our pumps and traps range from 400450µm, suggesting that they could have an extended competent stage.
Larval lifespan: 21 to >80 d, depending on temperature and food (see above)
Other attributes: Bathymodiolus is a broadcast spawner, producing large numbers of
offspring in a single spawning event Berg 198x.
Vestimentiferan (Siboglinid) Tubeworms
The giant siboglinid tubeworm Riftia pachyptila is a common,
habitat forming species at EPR vents. It grows quickly and
often displaces the related pioneer tubeworm colonist Tevnia
jerichonana in a successional sequence.
Size: ~ 0.1 mm for 34-d trochophore (Marsh et al. 2001), possibly ~0.2 mm at settlement
(this is the size of newly settled juvenile; Nussbaumer et al. 2006)
Sinking Velocity: Riftia eggs and zygotes ascend at 0.023 mm/s at deep-sea T and P
(Marsh et a. 2001); Capitella spp larvae (lecithotrophic) sink at 0.8-1.0 mm/s (Butman et
al. 1988)
Swim Speed: For a broad range of species, from both sessile and errant families, and in
stages from trochophore through nectochaete: 0. 8 to 5.2 mm/s (Konstantinova 1969
cited in Chia et al. 1984)
Precompetent stage length: Riftia trochophore larva, 34 d (as observed in Marsh et al.
2001). Riftia is a broadcast spawner, with a large brood size.
Feeding stage: Probably develops a mouth prior to settlement (Nussbaumer et al.
2006), but not known if it feeds in the plankton.
Competent stage length: If it does not feed in the plankton, 34 to 44 d total lifespan
(based on metabolic rates Marsh et al. 2001). The 34-d trochophore does not appear
competent to settle, suggesting the length of competent period is short. Perhaps not
more than about a week or two at 2°C (based on shallow water lecithotrophic
polychaete Capitella, which can delay metamorphosis for about 3 days at 15°C without
increased mortality). If it does feed, then possibly much longer
Estimated larval lifespan: 44 d maximum for nonfeeding larva, but could be longer if a
feeding stage develops.
Vent Polychaetes (e.g., Ophryotrocha, Amphisamytha)
Size: ~0.30 – 0.50 mm
Sinking Velocity: Lecithotrophic Capitella spp. larvae sink at 0.8-1.0 mm/s (Butman et
al. 1988)
Swim Speed: Polychaete larvae swim 0. 8 to 5.2 mm/s (Konstantinova 1969 cited in
Chia et al. 1984); 2-4 mm/s (Bolton and Havenhand 1997). Butman et al. 1988 found 2.53.3 mm/s on day 1 and 1.6-2.3 mm/s on day 3 for Capitella, which has no precompetency phase.
Precompetent stage length: Trochophore larva, probably lecithotrophic, as no feeding
larvae have been reported from the polychaete families to which they belong
(Dorvilleidae, Ampharetidae, Young, Larval Atlas, 2002)
Feeding stage: No feeding as larvae, but young juveniles are probably as mobile in the
plankton as the larvae. Some polynoids and terebellids are known to metamorphose in
the plankton and remain there as juveniles (“secondary larvae”, Young, Larval Atlas,
2002) for days to weeks in shallow water. Deep-sea species may do this too.
Competent stage length: Trochophores probably have a relatively short competent
phase, but the larvae are likely to be mobile for weeks after metamorphosis.
Larval lifespan: Non-feeding planktonic polychaetes in shallow water have typical
lifespans of 3 to 10 d at 10 to 15°C (Grantham et al. 2003).
References
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20 days (day 31 – 50) in competent stage
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behavior, morphology and metamorphic response. Mar. Ecol. Prog. Ser. 104:101108. (6 days in competent stage)
Hoegh-Guldberg, O., and J. S. Pearse. 1995. Temperature, food availability and the
development of marine invertebrate larvae. Am. Zool. 35:415–425.
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Metamorphic competence, a major adaptive convergence in marine invertebrate
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Pechenik, J.A. 1984. Relationship between rate of development and duration of larval
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