title used in an account of west-coast marine invertebrates entitled A Snail's Odyssey
  Predators & defenses
  Because defenses of adult barnacles are limited mainly to withdrawal into strong shells, this section on predators and defenses will focus on different predators. After some introductory material fishes as predators will be considered, while WHELKS, LIMPETS, SEA STARS, and INSECTS will be dealt with in other sections.
   
 

photograph in close view showing 1-2d-old spat of barnacles Chthamalus dalliMajor predators of larval acorn barnacles are juvenile fishes and a variety of suspension-feeding invertebrates, including adult barnacles.  Major predators of juvenile barnacles are limpets, whelks, and crabs, and major predators of adults are whelks and sea stars.  Defenses after metamorphosis include a strong outer calcified test and the ability to close up for long periods.

 



Spat of Chthamalus dalli settled onto the test plates of a
Balanus glandula.
The smallest individuals are likely just 1-2d old,
while some older C. dalli can be seen at the 7 o'clock position 8X

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Research study 1
 
No. 1 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli

The period just after metamorphosis is an especially risky time for all bottom-dwelling invertebrates, but moreso for sessile species such as barnacles.  Mortality for barnacles is highest during the first few moments of juvenile life, and may reach 40% or more in the first day.  By 1-2mo of age, up to 80% of the new settlers may be dead.  What are the causes of death? Let’s start with 100 newly settled Chthamalus dalli and consider some of the factors that will lead to their deaths over a 2-mo period.  Ideas from Gosselin & Qian 1997 Mar Ecol Progr Ser 146: 265 and Gosselin & Qian 1996 Mar Ecol Progr Ser 135: 69. 

NOTE the authors base their ideas on Balanus glandula recruits, but Chthamalus dalli recruits as shown here should work just as well

No. 2 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Predation   There are many animals including crabs, hermit crabs, worms, sea stars, and fishes that prowl the intertidal region looking for food.  The juveniles’ shell plates are too thin to offer much protection against these large predators. The X's mark a proportion of spat that are estimated will succumb to predators over the first 2mo of life.
No. 3 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Drying   Desiccation from air and/or sun exposure can be a serious cause of mortality in any small animal.  This partly owes to the unfavourable releationship of surface area to volume in small organisms.  Once a larger size is reached, drying effects are relatively less. More individuals in our array are X'd out.
No. 4 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Temperature extremes  These relate to tissue death apart from drying.  Hot and cold temperatures will have greater effect at higher positions in the intertidal zone. 
No. 5 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Sand abrasion/water motion  Both causes of mortality are obvious.  Shearing effects by waves and currents will have greater effect on smaller-sized individuals because of their greater relative surface areas
No. 6 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli UV radiation  Little is known about UV effects on invertebrates in temperate regions
No. 7 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Salinity  Low salinities caused by river discharge or localised sources of rainfall can seriously affect survival of many shore-inhabiting invertebrates. 
No. 8 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Energy depletion  Metamorphosis is metabolically demanding.  If insufficient energy stores are available to complete the process then the juvenile will die, or become stunted, perhaps too small and weak to withstand other environmental vicissitudes.
No. 9 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Bulldozing  Here, the scraping/dragging shells of hermit crabs, limpets, and other snails sweep off the metamorphosing larvae and early juveniles.
No. 10 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Space competition  This occurs as the juveniles get larger in size and, in barnacles, takes the form of overgrowing and undercutting by other individuals.
No. 11 in a series of photographs showing post-settlement mortality effects on a cohort of 100 barnacles Chthamalus dalli Food competition  This is less likely for a filter-feeding barnacle than for juveniles of motile species, like snails and worms, that have to forage for food. Nonetheless, a few individuals will die from this cause.
 
So, of the original 100 recruits to this area, only 20 have survived their first 2mo of life, as shown in this simulation on the Left. Many of the recruits have been eliminated completely, leaving fresh space on the rock for future colonisers, while others exist only as empty tests.
 
Research study 2
 

A study at Friday Harbor Laboratories, Washington shows that the effect of adult barnacles on consuming larval barnacles depends, in part, on species-specific behaviour of the larvae themselves.  For example, some species prefer to settle onto bare rock, while others favour the lateral shell-plates of the already existing large barnacles.  In some cases where adult Semibalanus cariosus densely populate the rock surfaces, a 65-100% reduction in barnacle settlement can be attributed to larval predation by adults.  Although most years are characterised by low survival of larvae, thus maintaining the bare patches, sometimes in a period of dense settlement of larvae recruitment of barnacles is high because their sheer numbers swamp the filtration ability photograph showing several barnacles Semibalanus cariosus on a rock with many other barnacles, most Balanus glandula, recruited onto their shell platesof the adult SemibalanusNavarrete & Wieters 2000 J Exp Mar Biol Ecol 253: 131.

NOTE  this is determined by the presence of nauplius larvae caught up in the cirri and present in the stomach contents of adult S. cariosus

NOTE  these newly recruited juveniles now have to face a gamut of hungry whelks and bulldozing limpets



Five adult barnacles Semibalanus cariosus have created an "oasis" of
settlement for other barnacles (mainly Balanus glandula) on
an otherwise bare rock surface. Note that many Balanus have
settled around the bases and opercular rims of their hosts, and
some even have survived to settle on the opercular plates 0.5X

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  Fishes
 
Research study 1
 

table showing relationship between density of juvenile rockfish in kelp beds and recruitment of barnacles Balalnus glandula in Monterey Bay, CaiforniaStudies in Monterey Bay, California on recruitment of barnacles Balanus glandula show a strong negative correlation of recruitment success with midsummer peaks in canopy area of offshore kelp beds Macrocystis pyrifera.  The reason is that juvenile rockfishes Sebastodes spp., known predators of barnacle nauplii and other invertebrate larvae, hang out under the canopy, presumably for their own protection.  In autumn and winter, predation on barnacle and other invertebrate larvae decreases owing to reduced densities of these planktivores.  The authors estimate that predation by the juvenile rockfish reduces recruitment of barnacles to 1/50th of the level that would occur in the predators' absence. Gaines & Roughgarden 1987 Science 235: 479.

NOTE measured as number of fishes seen per minute of SCUBA-diver observation

 
Research study 2
 

graph showing hiding times of barnacles Balanus glandula in relation to time spent feeding and after shadow-response stimulationgraph showing relationship between level of hunger in barnacles Balanus glandula and the length of time they hide after a shadow-responseShadow-responses, such as from predatory fishes passing overhead, are well known in marine invertebrates such as barnacles, tubeworms, and clams that rely on fast withdrawal into a shell, tube, or burrow to gain protection.  However, withdrawal may incur costs, such as oxygen deprivation or lack of feeding.  Studies in British Columbia show, for example, that barnacles Balanus glandula hide longer in response to a shadow passing overhead the more they have fed prior to the stimulus (see graph on Left). Size appears to have no effect on the hiding response.

graph showing the relationship of time to next air-exposure and duration of hiding time after a shadow stimulus in barnacles Balanus glandulaHiding times also decrease in proportion to hunger level (see graph upper Right).  In relation to this, however, the authors find that barnacles living higher on the shore (with less feeding opportunity) do not hide less than do barnacles living lower on the shore, where there is more time to feed.  Finally, what about the relationship between hiding time and time remaining to a receding tide when there is no opportunity to feed?  Would the barnacles exhibit a progressive decrease in hiding time? As seen in the graph on the lower Right, the answer is "yes'. All that remains now is to figure out how the barnacles do it. Dill & Gillett 1991 J Exp Mar Biol Ecol 153: 115.

NOTE  in this study, the time taken to resume fast cirral beating after a shadow is cast upon it

NOTE  here equated to starvation time

 
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