Mia J. Tegner

Kelp forest ecosystems: biodiversity, stability, resilience and future

Kelp forests are phyletically diverse, structurally complex and highly productive components of coldwater rocky marine coastlines. This paper reviews the conditions in which kelp forests develop globally and where, why and at what rate they become deforested. The ecology and long archaeological history of kelp forests are examined through case studies from southern California, the Aleutian Islands and the western North Atlantic, well-studied locations that represent the widest possible range in kelp forest biodiversity. Global distribution of kelp forests is physiologically constrained by light at high latitudes and by nutrients, warm temperatures and other macrophytes at low latitudes. Within mid-latitude belts (roughly 40–60° latitude in both hemispheres) well-developed kelp forests are most threatened by herbivory, usually from sea urchins. Overfishing and extirpation of highly valued vertebrate apex predators often triggered herbivore population increases, leading to widespread kelp deforestation. Such deforestations have the most profound and lasting impacts on species-depauperate systems, such as those in Alaska and the western North Atlantic. Globally urchin-induced deforestation has been increasing over the past 2–3 decades. Continued fishing down of coastal food webs has resulted in shifting harvesting targets from apex predators to their invertebrate prey, including kelp-grazing herbivores. The recent global expansion of sea urchin harvesting has led to the widespread extirpation of this herbivore, and kelp forests have returned in some locations but, for the first time, these forests are devoid of vertebrate apex predators. In the western North Atlantic, large predatory crabs have recently filled this void and they have become the new apex predator in this system. Similar shifts from fish- to crab-dominance may have occurred in coastal zones of the United Kingdom and Japan, where large predatory finfish were extirpated long ago. Three North American case studies of kelp forests were examined to determine their long history with humans and project the status of future kelp forests to the year 2025. Fishing impacts on kelp forest systems have been both profound and much longer in duration than previously thought. Archaeological data suggest that coastal peoples exploited kelp forest organisms for thousands of years, occasionally resulting in localized losses of apex predators, outbreaks of sea urchin populations and probably small-scale deforestation. Over the past two centuries, commercial exploitation for export led to the extirpation of sea urchin predators, such as the sea otter in the North Pacific and predatory fishes like the cod in the North Atlantic. The large-scale removal of predators for export markets increased sea urchin abundances and promoted the decline of kelp forests over vast areas. Despite southern California having one of the longest known associations with coastal kelp forests, widespread deforestation is rare. It is possible that functional redundancies among predators and herbivores make this most diverse system most stable. Such biodiverse kelp forests may also resist invasion from non-native species. In the species-depauperate western North Atlantic, introduced algal competitors carpet the benthos and threaten future kelp dominance. There, other non-native herbivores and predators have become established and dominant components of this system. Climate changes have had measurable impacts on kelp forest ecosystems and efforts to control the emission of greenhouse gasses should be a global priority. However, overfishing appears to be the greatest manageable threat to kelp forest ecosystems over the 2025 time horizon. Management should focus on minimizing fishing impacts and restoring populations of functionally important species in these systems.

Compounded Perturbations Yield Ecological Surprises

ABSTRACT All species have evolved in the presence of disturbance, and thus are in a sense matched to the recurrence pattern of the perturbations. Consequently, disturbances within the typical range, even at the extreme of that range as defined by large, infrequent disturbances (LIDs), usually result in little long-term change to the systems fundamental character. We argue that more serious ecological consequences result from compounded perturbations within the normative recovery time of the community in question. We consider both physically based disturbance (for example, storm, volcanic eruption, and forest fire) and biologically based disturbance of populations, such as overharvesting, invasion, and disease, and their interactions. Dispersal capability and measures of generation time or age to first reproduction of the species of interest seem to be the important metrics for scaling the size and frequency of disturbances among different types of ecosystems. We develop six scenarios that describe communities that have been subjected to multiple perturbations, either simultaneously or at a rate faster than the rate of recovery, and appear to have entered new domains or “ecological surprises.” In some cases, three or more disturbances seem to have been required to initiate the changed state. We argue that in a world of ever-more-pervasive anthropogenic impacts on natural communities coupled with the increasing certainty of global change, compounded perturbations and ecological surprises will become more common. Understanding these ecological synergisms will be basic to environmental management decisions of the 21st century.

Catastrophic Storms, El Niño, and Patch Stability in a Southern California Kelp Community

Strong winter storms in southern California destroyed most of the canopy ofthe giant kelp Macrocystis pyrifera but not the patches of understory kelps in the Point Loma kelp forest near San Diego. Subsequent massive recruitment of Macrocystis juveniles and adults that survived the storms had low survival in the summer during the California El Ni�o of 1983. The combined disturbance may have long-lasting structural consequences for this community because, once established, the understory patches can resist invasion by Macrocystis.

No-take Reserve Networks: Sustaining Fishery Populations and Marine Ecosystems

Abstract Improved management approaches are needed to reduce the rate at which humans are depleting exploited marine populations and degrading marine ecosystems. Networks of no-take marine reserves are promising management tools because of their potential to (1) protect coastal ecosystem structure and functioning, (2) benefit exploited populations and fisheries, (3) improve scientific understanding of marine ecosystems, and (4) provide enriched opportunities for non-extractive human activities. By protecting marine ecosystems and their populations, no-take reserve networks can reduce risk by providing important insurance for fishery managers against overexploitation of individual populations. Replicated reserves also foster strong scientific testing of fishery and conservation management strategies. Reserve networks will require social acceptance, adequate enforcement, and effective scientific evaluation to be successful. Processes for reserve establishment should accommodate adaptive management so bounda...

El Niño Effects on Southern California Kelp Forest Communities

Publisher Summary The chapter reviews the effects of the massive oceanic phenomenon on the nearshore kelp communities of the southern California Bight. The effects of this El Nino on intertidal algae in southern California are ambiguous; Gunnill is unable to find a uniform response in seven species of macro-algae. The effects of the 1982-1983 El Nino on California kelp forest communities varied both locally and regionally. The relevance of El Ninos to kelp forest community structure must be related to recurrence rates. The effects on kelp community structure will depend on the season of the warm period for example, given the normal summer stress on Macrocystis, warm-water events then are more serious than a similar temperature elevation in winter, and the duration and magnitude of warming. Two major El Ninos stand out in recent California history, the events of 1957-1959 and 1982-1984 from the biological perspective, the opposite side of the coin may be even more important to the California Current system. Anti El Nino years, characterized by strong southward transport, low sea level, reduced temperature and salinity, and high zooplankton abundance, are years of high biological productivity.

Protease released from sea urchin eggs at fertilization alters the vitelline layer and aids in preventing polyspermy

Abstract Sea urchin eggs release protease activity extracellularly from 30 to about 60 sec after insemination. This timing corresponds to the breakdown of the cortical granules. The evidence presented shows that the protease prevents polyspermy through its action on the vitelline layer. The protease ‘hardens’ the layer and makes it incapable of binding sperm. Sperm that are bound to the vitelline layer before cortical granule breakdown lose their attachment to the layer as the protease is excreted. By digesting the bonds between supernumerary sperm and the vitelline layer and by ‘hardening’ the layer the protease aids in establishing the block against polyspermy.

Storm wave induced mortality of giant kelp, Macrocystis pyrifera, in Southern California

orbital velocities (associated with large, high frequency waves), the presence of breaking waves, and entanglement by drifters were found to increase mortality through stipe breakage or holdfast failure. Longshore variability in wave intensity was found to affect kelp mortality rates. The data suggest that wave breaking may be an important factor in determining the inner boundary of the kelp bed.

Over-exploitation of a broadcast spawning marine invertebrate: Decline ofthe white abalone

Marine invertebrates have long been consideredto be resistant to overfishing. However, agrowing number of exploited taxa have declinedsubstantially and even disappeared from partsof their former range. We consider the case ofthe white abalone (Haliotis sorenseni);the first marine invertebrate proposed for theUS endangered species list. This high-valuespecies was one of five abalones targeted inthe California and Mexico fisheries; it is nowrare and protected from fishing. The biologicalcharacteristics of this deep-living abaloneindicate that it was particularly vulnerable toover-exploitation; reduction of density orgroup size is now known to lead to declines infertilization success and recruitment failure.Warning signs of potential problems existedboth pre- and post-exploitation but were notrecognized. In particular, serial depletion wasnot detected because catch was not analyzedspatially, perhaps because total landings werereasonably stable for the short period ofexploitation. Recent submersible surveys led toestimates that white abalone now number lessthan 2,600 animals or 0.1% of the estimatedpre-exploitation population size. Densities andestimated population sizes are less than 100animals, at all but one location. Alternateexplanations for the decline in abundance wereconsidered and only exploitation-linkedfactors, such as sub-legal mortality andillegal fishing, were likely contributors.Episodic recruitment appears to be acharacteristic of broadcast-spawning,long-lived species and may make themparticularly vulnerable to over-exploitation.Management strategies based on size limits thatallow a few years of spawning prior to reachingminimum legal size are insufficient.Sustainable fisheries will require multipleprotected areas to preserve brood stockaggregations necessary for successfulfertilization.

OCEAN WARMING EFFECTS ON GROWTH, REPRODUCTION, AND SURVIVORSHIP OF SOUTHERN CALIFORNIA ABALONE

Traditional fisheries management in southern California has failed, in part because it is based on an assumption of an unvarying environment and is focused on size limits rather than insuring the persistence of aggregations of large fecund individuals. The combined effect of low frequency climatic variability and anthropogenic perturbations can have dramatic consequences for abalone in southern California. Abalone species are tightly linked to kelp forest ecosystems that, besides furnishing habitat, also provide the main food source for abalone. In southern California, kelp canopies are very sensitive to oceanographic climate because the kelp depend upon high nutrients in the water column. Oceanic warming, in turn, results in decreased nutrients in the surface water, and this is correlated with marked reductions in giant kelp biomass. Here we address the additive effects of ocean warming on two species of California abalone (the red abalone, Haliotis rufescens; and the green abalone, H. fulgens) by subjecting them to varied environmental conditions similar to cool, normal, and warm phases of the California current in the southern California Bight. Our experimental design simultaneously tested the synergistic effects of temperature and food quantity and quality on survivorship, growth, and reproduction. For red abalone, warm temperatures increased the onset of with- ering syndrome, a fatal abalone disease, and halted growth and reproduction. In contrast, green abalone survivorship, growth, and reproduction were relatively robust irrespective of temperature, while their growth and reproduction were most strongly influenced by food quantity. We found clear evidence suggesting that, combined with overfishing, California abalone populations are adversely affected by ecosystem responses to ocean warming: Cool- water red abalone suffer stronger consequences in warm water than do green abalone. Conservation, restoration, and recovery plans of remnant California abalone populations

Sea Urchin Sperm-Egg Interactions Studied with the Scanning Electron Microscope

Scanning electron microscopy of the outer surface of sea urchin eggs sampled at intervals during the first 3 minutes after insemination reveals the detailed structural changes of the vitelline layer during its transformation into the fertilization membrane. A sperm attachment-detachment sequence is described for the large number of sperm which transitorily bind every egg during fertilization.