Research Notes: Mobility, Gregariousness and Attachment in Four Small Bivalve Mollusc Species at Husvik, South Georgia
Research Notes: Mobility, Gregariousness and Attachment in Four Small Bivalve Mollusc Species at Husvik, South Georgia
Research Notes: Mobility, Gregariousness and Attachment in Four Small Bivalve Mollusc Species at Husvik, South Georgia
RESEARCH NOTES
Mobility, gregariousness and attachment in four small bivalve mollusc species at
Husvik, South Georgia
J. Davenport and P.C. Wilson
University Marine Biological Station, Millport, Isle of Cumbrae, Scotland, KA28 OEG, U.K.
Figure 2. Byssus of Lissarca miliaris. A. Multiple threads (arrowed) radiating from byssus opening onto
surface of piece of frond of Schizoseris (scale bar = 100 u,m). Bivalve (b) is at bottom of photograph, fragments
of alga (a) are at top. B. Closeup of slightly triangular attachment (c) of byssus thread (arrowed) to sea weed
(scale bar = 50 jun).
Downloaded from http://mollus.oxfordjournals.org/ at University of Saskatchewan on September 19, 2012
Kit
RESEARCH NOTES 495
Figure 3. Byssus of Lasaea rubra. A. View of two animals connected by byssus threads (scale bar = 500 u,m).
B. Closeup of byssus thread (arrowed) and attachment (a) to shell of conspecific (scale bar = 10 UJTI).
vations were made at Husvik under a binocular mobility largely ceased once a group had been
microscope. formed. Kidderia also formed numerous attachments
Equipment was not available to conduct a thorough to the substratum, as well as to conspecifics.
assessment4 of the mechanical properties of byssus. Table 3 shows speed of movement in the four
Instead, a crude estimate of byssus elasticity was species. The two species associated with seaweeds
obtained for 5 animals of each species in the following (Gaimardia and Lissarca) have similar relative speeds
manner. Each animal was left in a petri dish of sea- (shell length s"1) and move much more quickly (both
water for one hour so that it attached either to the absolutely and relatively) than the two species that
dish or to another animal. Beneath a binocular micro- attach to rock (Kidderia and Lasaea). Kidderia moves
scope, all byssus threads except one were cut. The much more slowly than does Lasaea, confirming the
Caimardia trapesina 20 0
Lissarca miliaris 18 2
Lasaea rubra 0 20
Kidderia bicolor 0 20
moving water if they were not attached by byssus. the weed. Second, recovery from dislodgement is
Gaimardia and Lissarca were both characterized by virtually impossible; Macrocystis, though attached to
an unusual reaction to touch. Most bivalve molluscs the substratum by a massive holdfast, is essentially a
(including Lasaea and Kidderia) respond to touch by tethered surface weed and a bivalve that falls from it
immediate foot retraction and shell valve adduction. is most unlikely to be able to regain contact with kelp.
The two weed-dwellers barely responded to a light A highly streamlined shell and strong, elastic byssus
touch, while stronger tactile stimulation provoked counteract these problems. Because the byssus thread
brief shell valve adduction, but not foot retraction. is very long, the animal can move rapidly around on
Only prolonged stimulation could invoke a 'normal' the weed without losing a firm attachment. During
closure response and then only for a few seconds. periods when the animal detaches the byssus the
Garimardia and Lissarca both secreted byssus very unusually strong pedal adhesion provides further pro-
rapidly (often in less than 60s) when taken from sea- tection. The overriding necessity of avoiding dis-
weed and placed in holding vessels. Attachment time lodgement presumably explains the suppression of
in the other two species was very variable and much the normal bivalve closure response to touch.
slower; several minutes often elapsed before byssus Lissarca faces similar problems, particularly when
secretion. living on intertidal Schizoseris. This complex red alga
Observations in early April confirmed the pre- is swept to and fro on every wave at the time of low
carious nature of the lifestyle of Gaimardia and water, presumably causing oscillations in inertial
Lissarca. After the storm of 28-31st March, 1994, forces acting upon the Lissarca. A highly elastic
large numbers of both species were found either dead byssus will help to absorb these shocks. Lissarca often
on the shore or living on the 'wrong' substrata. Several occurs on bare rock around Schizoseris plants, es-
Gaimardia were seen attached to middle shore pecially in surge channels. It seems probable that
Porphyra or lower shore Palmaria, specimens of Lissarca are sometimes dislodged from Schizoseris —
Lissarca were also seen on Porphyra, or on bare rock, they may well be able to regain their position by
but there were many floating in the surf, their shells virtue of a strongly adhesive foot and high level of
filled with air. In contrast, there was no evidence of mobility. The suppression of the normal shell valve
dead or dying Kidderia or Lasaea. closure response in both Gaimardia and Lissarca is an
The four species show a diversity of approaches to interesting example of convergence, given their
solving the problem of attachment in a high energy taxonomic separation.
environment. Gaimardia, living on Macrocyslis, faces Kidderia often lives on bare rock, battered by
two main problems. First, the strong wind-generated waves. It is unlikely to encounter the inertial prob-
surface currents and exposure to oceanic swell mean lems associated with being swept to and fro in weed
that the animals are exposed to considerable and - when living amongst Iridea carpet it lives amongst
highly variable forces tending to dislodge them from the matted holdfasts, not the fronds. Kidderia clearly
498 RESEARCH NOTES
does not need highly elastic byssus; instead it requires
1
a tightly bound, firm attachment; it also relies on
gregarious attachment, often with thousands of ani-
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(shell len
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dodo microhabitat for newly-released brooded young.
Rapid movement is unnecessary in this species.
Lasaea rubra lives in a more protected environ-
ment, avoids exposed rock or plant surfaces. The
lower risk of dislodgement permits lower pedal ad-
hesion, a relatively weak and fine byssus, yet a high
7 level of mobility (allowing invasion of new microhabi-
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CD 0 j j One of the authors (JD) acknowledges the financial
i REFERENCES
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i- p p p
1. Ralph, R. & Everson, I. 1972. Brit. Ant. Surv.
CO II Bull., 31: 51-54.
1 d d d d
— CO
2. RICHARDSON, M.G. 1979. Brit. AM. Surv. Bull.,
E 49: 91-115.
3. COOLEY, W.A., SCOTT, A.C., & SIMMONS, M.M.
1994. Microsc. & Analysis, 43: 23-25.
4. SMEATHERS, J.F. & VINCENT, J.F.V. 1979. J. Moll.
Stud., 45: 219-230.
I
c
CD O 5. DENNY, M.W. 1988. Biology and the mechanics of
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i^ in co co the wave-swept environment. Princeton University
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