The association of coloniality with parental care of embryos

Many colonial marine animals care for embryos by brooding them on or in their bodies. For brooding to occur, features of the animals must allow it, and brooding must be at least as advantageous as releasing gametes or zygotes. Shared features of diverse colonial brooders are suspension feeding and a body composed of small modules that are indefinitely repeated and can function semi-autonomously, such as polyps or zooids. Suspension feeding permits capture of sperm for fertilization of ova that are retained by the parent. Distribution of broods among numerous small polyps, zooids, or other small modules facilitates supply of oxygen to embryos that are retained and protected by the parent. Brooding increases survival of offspring, controls dispersal, and can provide other developmental advantages. Colonial ascidians, pterobranch hemichordates, and entoprocts brood; most bryozoans and many colonial cnidarians brood. An unanswered question is why so many colonial anthozoans do not brood. Sponges share with colonies capacities for capturing sperm and separating numerous retained embryos yet many do not brood. Hypotheses for nonbrooding by colonies and sponges necessarily must apply to particular taxa. Few have been tested.