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Reproduction |
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Settlement & metamorphosis |
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Settlement & metamorphosis are dealt with in this section, while topics of
MATE SELECTION & COPULATION,
EGG-LAYING,
EMBRYONIC DEVELOPMENT,
HATCHING & LARVAL DEVELOPMENT,
SETTLEMENT CUES, and
ONTOGENETIC DEVELOPMENT OF BEHAVIOUR are considered elsewhere. |
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Research study 1 |
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In what is thought to be the first time a marine gastropod with long-duration planktotrophic development has been successfully and reliably cultured, researchers at the Marine Biological Laboratory, Massachusetts rear Aplysia californica from egg through metamorphosis. After 34d of larval development on a diet of unicellular alga Isochrysis galbana (22oC), and stimulated by the presence of the red alga Laurencia pacifica, the veligers settle and metamorphose. Within 3-4d after the onset of metamorphosis the juveniles begin feeding on Laurencia and other red seaweeds and, by 120d after hatching and at a mass of about 400g, are sexually mature. Of 6 species of west-coast algae tested as metamorphosis inducers, only Laurencia pacifica is successful. Development of rearing techniques for A. californica has been a truly ground-breaking achievement and has enabled research specimens for neurophysiological research to be cultured essentially ad libitum. Kriegstein et al. 1974 Proc Nat Acad Sci, USA 71: 3654.
NOTE the authors note that 4 individuals of A. californica were previously cultured to reproductive maturity at the Pacific Bio-Marine Supply Co., California but that the techniques did not prove reliable
NOTE a culture facility, the National Resource for Aplysia, now exists on the campus of the University of Miami’s Rosenstiel School of Marine & Atmospheric Science, and provides annually about 30,000 sea hares of all ages and sizes to neuroscience researchers around the world over |
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Research study 2 |
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During metamorphosis in nudibranchs there is, as in most other marine invertebrates, a considerable re-arranging of tissues and structures. An individual changes within a day or so from a free-swimming, phytoplankton-eating, shelled organism to a crawling, predatory, shell-less form. All soft tissues are resorbed and used; only the larval shell is cast off. Drawings modified from Bonar 1978 p. 177 In, Settlement and metamorphosis of marine invertebrate larvae (Chia & Rice, eds.) Elsevier, N.Y. |
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Research study 3 |
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A post-metamorphic 5mm-long Dirona albolineata is already feeding on adult-type foods (snails, bryozoans, hydroids). Drawing from Hurst 1967 Veliger 9: 255.
Without the oral veil spread out on its anterior
end, it is difficult to tell which end is which in
this adult Dirona albolineata 1X
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Research study 4 |
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Veligers of Hermissenda crassicornis settle from the plankton after 5wk in response to chemical emanations from hydroid prey such as Obelia spp. Metamorphosis is completed within 2-3d and within a few days the juvenile, at 0.4mm in length, begins to feed on hydroids. Hermissenda’s complete lifespan is about 4mo. Harrigan & Alkon 1978 Biol Bull 154: 430.
NOTE at 15°C in laboratory culture
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Research study 5 |
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The subtidal nudibranch Tritonia diomedea can be readily cultured in the laboratory. Development at 12°C leads to completion of metamorphosis within 34d. Metamorphosis will take place in the absence of a specific substratum. Kempf & Willows 1977 J Exp Mar Biol Ecol 30: 261. Photograph courtesy Russ Wyeth & Owen Woodward. |
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Research study 6 |
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 Studies on larval development of specimens of Melibe leonina collected in the San Juan Archipelago and southern Vancouver Island regions show a similar pattern to that described for other species of opisthobranchs that have planktotrophic larvae. The graph on the Right shows 3 stages of larval development (see blue dots): hatching (Day 0), eyespot-mantle retraction (Day 16-20 post-hatching), and commencement of metamorphosis (30-48d post-hatching).
The rudiments of cerata and oral hood appear in the late-stage larva. Metamorphosis involves loss of shell, operculum, and velum. Some parts of the larval stomach are retained along with right and left digestive diverticula. The photo-series at the bottom shows metamorphic events during the 48-60h of metamorphosis (12-14°C). Shortly after metamorphosis there is a rapid expansion of the primary cerata and oral hood.
Within 2-3d the juvenile is using its oral hood to catch prey (see figure on Left). Juveniles can be maintained on a diet of ciliates and copepod nauplii. The authors note that a specific, external chemical cue does not appear to be required for metamorphosis. Bickell & Kempf 1983 Biol Bull 165: 119. Colour photo of Melibe leonina courtesy Charles Seabourne, Los Angeles.
NOTE at this time the epithelium of the mantle fold detaches from the rim of the shell and is pulled posteriorly. The tissues coalesce and eventually grow into two protuberances – these are the rudiments of the primary cerata

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Research study 7 |
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Settlement of larval sea hares Aplysia californica on rocky reefs off the coast of Santa Catalina Island, California is primarily to the red alga Plocamium cartilagineum. There are 2 settlement pulses, one in Jan-Feb probably from local spawnings the previous summer, and one in Jun-Sept possibly from larvae spawned in the region of Baja California and carried northwards in summercurrents. Individuals from the winter recruitment period are 500g or more in size by the time they reproduce in summer, while ones from the summer recruitment are only about 1g when they reproduce in summer. Pennings 1991 J Exp Mar Biol Ecol 146: 253. Photograph courtesy Kevin Lee, Fullerton, California diverkevin.
Juvenile sea hare Aplysia californica of just a few cm
in size. Note the much smaller opisthobranch of uncertain
ID crawlingin the reverse direction on the frond 2X |
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Research study 8 |
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The transient nature of nudibranch recruitment to our shores is familiar to marine biologists – sometimes they’re there, sometimes not. Clearly, some link in the larval production/survival/transport chain is involved, but what it is, and how and when it works has not been identified. Recently, sharp declines in numbers of nudibranchs in central California have been correlated with La Niña climatic events that disrupt onshore and alongshore currents that would otherwise bring larvae into favourable areas for settlement and metamophosis. To investigate this, a research group from a variety of Californian and other institutions model distribution and abundance data for 56 nudibranch species collected between 1969-1995 with such ocean-climate variables as sea-surface temperature, sea-surface height, sea-level atmospheric pressure, and upwelling strength, to predict most favourable conditions for larval settlement. The results of this wonderfully ambitious and broad-scale study support a cross-shelf and alongshore larval-advection theory to explain past and current fluctuations in abundances of nudibranchs in central California. When sea levels are high and inshore cross-shelf currents predominate, as during El Niño conditions, recruitment is high; during the opposite conditions of La Niña events, inshore movements of larvae are not favoured, and recruitment is low (by up to 2 orders of magnitude). Given that the latter conditions have predominated for several years, the authors hypothesise that nudibranch populations will likely recover when El Niño conditions return. Schultz et al. 2011 Limnol Oceanogr 56: 749. Photograph courtesy Kevin Lee, Fullerton, CA.
NOTE major hypotheses involve changes in sea-surface temperature, nutrient levels for sustenance of larvae, and current direction and strength (cross-shelf larval advection)
NOTE the normal summer flow of the California Current is southward and offshore, which increases coastal upwelling. El Niño events transiently reverse the current to northward and onshore, which produces downwelling
More than one or two opisthobranchs being found together at
one time is usually rare. Perhaps the presence of several
dozens of the cephalaspidean Bulla gouldiana in this southern
California site has resulted from a favourable El Nino event 0.2X
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