INTRODUCTION ... . .. ..... .. . ..... ...... ..... .. 195 ALGAE 197 LICHENS .. ....... ...... .... .... . ....... ...... ..... ..... . 199 FUNGI ...... .... ....... ...... .... 204 BRYOPHYTES, PTERIDOPHYTES, AND SPERMATOPHYTES 205 RNA and Protein Synthesis 206 Mosses •......• 206 Seeds 208 Des��cation. oj Germinating .Se�ds and Seedlings 211 'Presowmg drought hardenmg 212 Photosynthesis, Respiration, and Dark Fixation oj CO} 2 1 3 Bryophytes 2 1 3 Vegetative tissues o f vascular plants 2 1 7 Seeds 218 Ultrastructure, Permeability, and Membranes, and the Possible Bases of Desiccation Tolerance 220

Physiological Aspects of Desiccation Tolerance

Seaweeds are ecologically important primary producers, competitors, and ecosystem engineers that play a central role in coastal habitats ranging from kelp forests to coral reefs. Although seaweeds are known to be vulnerable to physical and chemical changes in the marine environment, the impacts of ongoing and future anthropogenic climate change in seaweed-dominated ecosystems remain poorly understood. In this review, we describe the ways in which changes in the environment directly affect seaweeds in terms of their physiology, growth, reproduction, and survival. We consider the extent to which seaweed species may be able to respond to these changes via adaptation or migration. We also examine the extensive reshuffling of communities that is occurring as the ecological balance between competing species changes, and as top-down control by herbivores becomes stronger or weaker. Finally, we delve into some of the ecosystem-level responses to these changes, including changes in primary productivity, diversity, and resilience. Although there are several key areas in which ecological insight is lacking, we suggest that reasonable climate-related hypotheses can be developed and tested based on current information. By strategically prioritizing research in the areas of complex environmental variation, multiple stressor effects, evolutionary adaptation, and population, community, and ecosystem-level responses, we can rapidly build upon our current understanding of seaweed biology and climate change ecology to more effectively conserve and manage coastal ecosystems.

Effects of climate change on global seaweed communities

Rocky shores occur at the interface of the land and sea. Typically they are open ecosystems, with steep environmental gradients. Their accessibility to man has rendered them susceptible to a variety of impacts since prehistoric times. Access can be regulated, however, and they are more amenable to management than open ocean habitats. This review uses examples from throughout the world to demonstrate the extent to which rocky shores have been, and are currently, affected by pollution (examples used are endocrine disrupters, oil, eutrophication), over-collection of living resources, introduced alien species, modification of coastal processes (coastal defences, siltation) and global change (climate, sea level). These impacts are put into the context of natural fluctuations in time and variability in space of both the environment and the organisms. The relative magnitudes of some anthropogenic disturbances differ between the industrialized, developed world and the developing world. For example, in developed, industrialized countries pollution based impacts should diminish over the next 25 years due to improved regulation and a reduction in older ‘dirtier’ heavy industry. Conversely, in many developing countries pollution will increase as a consequence of growth in the human population and industrialization. Except for large-scale disasters such as oil spills, pollution tends mainly to influence embayed coastlines. Chronic effects such as eutrophication can have broader-scale impacts over whole coastlines and elevated nutrient levels have also been implicated in a trend of increasing frequency of catastrophic kills due to harmful algal. Direct removal of living resources has had major effects on coastlines at both local and regional scales and is likely to increase over the next 25 years, especially in developing countries where rapidly expanding human populations will put further pressure on resources. Impacts from recreational activities are likely to increase with greater leisure time in wealthier regions of the world, and cheaper travel will spread these impacts to poorer regions. Invasions by alien species have increased in frequency during the last 20 years leading to some dramatic effects on native assemblages. Problems associated with alien species, especially pathogens, will continue to increase over the next few decades. The proportion of the coastline modified by artificial structures (breakwaters, seawalls, groynes) will increase because of coastal development and defences against sea-level rise and the greater frequency of storms. This will increase connectivity between areas of rocky habitat. Siltation will continue to increase due to urbanization of catchments and estuaries, and changes in agricultural practice. This may have considerable impacts at local and regional scales, favouring sediment tolerant organisms such as turf algae and anemones. In the future, greater frequency of environmental extremes is likely, including large-scale events such as the El Nino Southern Oscillation (ENSO). Global change in temperature, sea-level rise and increases in the frequency of storms will affect rocky shores throughout the world, but this will occur over long time scales; over the next 25 years most of the responses by rocky shore communities will mostly be quite subtle. Thus rocky shores will be subject to increasing degradation over the next 25 years. They are, however, less vulnerable than many other aquatic habitats due to their hard substratum (rock), their relative lack of large biogenic structures and to their generally open nature. They are also remarkably resilient, and recovery can occur rapidly due to recruitment from unaffected areas. Their susceptibility to both terrestrial and marine disturbances does make them more vulnerable than sublittoral and offshore habitats. There are considerable gaps in knowledge, particularly of certain microhabitats such as crevices, boulders, sand-scoured areas and rock pools. These have been much less studied than more accessible assemblages on open, freely draining rock. More research is needed to establish the effects of increasing sediment loads, ultraviolet radiation and introduced species on rocky shore communities. Strategic and applied research programmes should integrate field experiments and carefully selected monitoring programmes to verify management regimes. Hindcasting from the palaeo-record would be valuable, to compare rates of predicted change with periods when change was rapid in the past. This information could, in principle, be used to help conserve rocky shores through networks of marine protected areas and a general reduction of environmental pollution.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0376892902000115

Rocky intertidal communities: past environmental changes, present status and predictions for the next 25 years

Intertidal seaweeds are periodically exposed to air where they experience a variety of potentially stressful environmental conditions, including nutrient limitation, high light, high and low temperature, desiccation, and osmotic stress. This paper considers the current understanding of stress tolerance in intertidal seaweeds and discusses ways in which future research could increase our understanding of the role of environmental factors in the ecology and physiology of these algae. We believe research is required in at least three areas. 1) Laboratory physiological studies have established that correlations exist between stress tolerance and the vertical distribution of species. However, little information is available on the importance of stress in determining community structure in nature. Field experiments are essential to relate the impact of single or multiple stresses on the survival, growth, and reproductive output of macroalgae. In paticular, it is necessary to clarify the role of sublethal stress in determining the outcome of competitive interactions. 2) With the exception of obvious lethal effects or damage associated with extreme environmental conditions, such as unusually hot and dry weather, it is difficult to evaluate the occurrence and severity of stress in natural populations of seaweeds. There is a need to develop molecular and biochemical markers specific for individual stresses or groups of stresses to allow the unambiguous and direct determination of stress in situ. 3) Despite the apparent importance of stress in intertidal seaweeds, we are largely ignorant of the mechanistic basis of tolerance. The application of currently available tools of molecular and cell biology to the investigation of stress-induced transcriptional and translational changes could enormously increase our understanding of both the sites of, and pathways involved in, stress tolerance. In summary, there are numerous unanswered fundamental questions about the stress tolerance of intertidal seaweeds, providing opportunities for research ranging from field ecology to molecular biology and biochemistry.

Stress tolerance in intertidal seaweeds

The structure of a tropical rocky intertidal community on Taboguilla Island on the Pacific coast of Panama is characterized by extremely low abundances of noncrustose algae and sessile animals, indistinct vertical zonation patterns (a result of the low abundances), and the occurrence of most invertebrates (except barnacles) and upright algae in holes and crevices (as opposed to open, smooth surfaces). This contrasts strikingly with two temperate rocky intertidal communities, which have high covers of plants and animals, more obvious zones, and invertebrates and upright algae occurring both on relatively homogeneous surfaces and in holes and crevices. Field experiments were done and observations were made in the Panama community to test the effects of different types of consumers (both predators and herbivores) on their prey and on the types of escapes utilized by the prey. Consumer exclusion experiments suggest that (1) predation and herbivory are severe at all times of the year, (2) consumer pressure is a cumulative function of many types and species of predators and herbivores, (3) the primary effect of larger fishes and crabs is to restrict most prey to three—dimensional refuges (holes and crevices), and (4) the primary effect of smaller consumers, mostly invertebrates, is to keep abundances of the prey low. Thus, in the Panama system, three—dimensional space (holes and crevices) appears to be particularly important as a refuge from consumers, while escapes from consumers in body size, time, or two—dimensional space (e.g., in a higher zone) documented so frequently in temperate areas, assume secondary importance for many prey. This restriction of the types of escapes utilized by prey species appears to be a consequence of two main factors: the presence of fast—moving consumers (i.e., herbivorous and predaceous fishes and herbivorous crabs which are absent or rare in the two temperate communities), and the year—round foraging of all consumers.

Mark W. Schonbeck

Papers

Factors controlling the lower limits of fucoid algae on the shore

The effects of brief periodic submergence on intertidal fucoid algae