Succession and Stability Outline Concepts 456 » 20-1 Community Changes During Succession 456 COVikipl 20.1 Review 460 "20.2 Ecosystem Changes During Succession 460 Coricept 20.2 Review 465 \ \2h.seMech@nismsof Succession 465 Concep 20.3 Review 469 204 ‘Community and Ecosystem Stability 469 __ Investigating the Evidence 20: Variation Around the Median 472 “ Concept 20.4 Review “474 “Applications: Using Repeat Photography to Detect Suz Long-Term Change 474 ‘Summary 477 i. Key Terms 477 Review Questions 477 Suggested Readings 478Te first recorded visit to Glacier Bay gave no hint of its eventual contributions to our understanding of biological communities and ecosystems. In 1794. Captain George Vancouver visited the inlet to what is today called Glacier Bay, Alaska (lig. 20.1). He could not pass beyond the inlet to the bay. however, because his way was blocked by a mountain of ice. Vancouver and Vancouver (1798) described the scene as follows: “The shores of the con- ‘inent form two large open bays which were terminated by ‘compact solid mountains of ice, rising perpendicularly from the water's edge, and bounded to the north by a continuation of the united lofiy frozen mountains that extend exstward from Mount Fairweather. In 1879, John Muir explored the coast of Alas} relying heavily on Vancouver's earlier descriptions, Muir (1915) commented in his journal that Vancouver's descrip- tions were excellent guides except for the area within Glacier Bay. Where Vancouver had met “mountains of ice,” Muir found open water. He and his guides from the Hoona tribe paddled their cance through Glacier Bay in rain and mist, feeling their way through uncharted territory. They eventually found the glaciers, which Muir estimated had retreated 30 to 40 km up the glacial valley since Vancouver's isit 85 years earlier. Figure 20.1 Glacier Bay. Alaska, Chapter 20. Succession and Stability 455° ‘Muir found no forests at the upper portions of the bay. He ‘and his party had to build their campfires with the stumps and trunks of long-dead trees exposed by the retreating glaciers. Muir recognized that this “fossil wood” was a remnant of a forest that had been covered by advancing glaciers centuries earlier. He also saw that plants quickly colonized the areas uncovered by glaciers and that the oldest exposed areas, ‘where Vancouver had met his mountains of ice. already sup- ported forests. Muir's observations in Glacier Bay were published in 1915 and read the same year by the ecologist William S. Cooper. Encouraged by Muir’s descriptions. Cooper visited Glacier Bay in 1916 in what was the beginning of a lifetime of study (Cooper 1923, 1931, 1939). Cooper saw Glacier Bay as the ideal laboratory for the study of ecological succession, the change in plant, animal, and microbial com- ‘munities in an area following disturbance or the creation of new substrate, Glacier Bay was ideal for the study of sue~ cession because the history of glacial retreat could be accu- rately traced back to 1794 and pethaps farther. Cooper ultimately made four expeditions to Glacier Bay. His work and that of later ecologists produced a detailed picture of succession there. Several species of plants colonize an area during the first 20 years after it is456 Section V exposed by the retreating glacier. These plants, the first in a successional sequence. form a pioneer community. The most common members ofthe pioneer community are horse tail. Equiserum varietaum. willow herb, Epilobium tati- folium. willows, Salix sp.. cottonwood seedlings, Populus trichocarpa, mountain avens, Dryas drummondii, and Sitka spruce, Picea sitchensis. ‘About 30 years after an area is exposed, the pioneer community gradually grades into a community dominated by mats of Dryas, a dwarf shrub. These Dryas mats also contain scattered alder, Alnus sinuata, Salix, Populus, and Picea. Then, about 40 years after glacial retreat, the community changes into a shrub-thicket dominated by Alnus. Soon after the closure of the Alnus thicket, however, Populus and Picea will grow above it, covering about 50% of the area on sites 50 to 70 years old. In 75 to 100 years, succession leads to a forest commu- nity dominated by Picea, Mosses carpet the understory of this spruce forest and here and there grow seedlings of western hemlock, Tiuga heterophylla, and mountain hemlock, Tstuga ‘mertensiana. Eventually. the population of Picea declines and the forests are dominated by Tsuga. On landscapes with shallow slopes these hemlock forests eventually give way to muskeg, a landscape of peat bogs and scattered tussock meadows. Because succession around Glacier Bay occurs on newly exposed geological substrates, not significantly mod- ified by organisms, ecologists refer to this process as primary suecession, Primary succession also occurs on newly formed volcanic surfaces such as lava flows. In areas where disturbance destroys a community without destroying the soil, the subsequent succession is called secondary succes- jon. For instance, secondary succession occurs after agri- ‘cultural lands are abandoned or after a forest fire. Succession generally ends with a community whose populations remain stable until disrupted by disturbance. ‘This late successional community is called the climax com- munity. The nature of the climax community depends upon environmental circumstances. The communities we dis- cussed in chapter 2—temperate forests, grasslands, ete.— ‘were essentially the climax communities for each of the climatic regimes that we considered. The nity around Glacier Bay is determined by the prevailing cli mate and local topography. On well-drained, steep slopes the climax community is hemlock forest. In poorly drained soil on shallow slopes the climax community is muskeg. Studies of succession show that communities and ecosystems are not static but constantly change in response to disturbance, environmental change, and their own internal dynamics. In many cases, the general direction of change in community structure and ecosystem processes is. pre- dictable, atleast over the short term. The patterns of change in community and ecosystem properties during succession and the mechanisms responsible for those changes are sub- |jects covered in chapter 20. We also consider a companion topic, community and ecosystem stability. Communities and Ecosystems mot. as 20.1 Community changes during succession include increases in species diversity and changesin species composition. ‘ 20.2 Ecosystem changes during succession include increases in biomass, primary production, respiration, and nutrient retention. 20.3 Mechanisms that drive ecological succession include facilitation, tolerance, and inhibition. 20.4 Community stability may be due to lack of disturbance or community resistance or resilience the face of disturbance. Community Changes During Succession Community changes during suecession include increases in species diversity and changes in species composition. Some of the most detailed studies of ecological succession have focused on succession leading to a forest climax. ‘Though primary and secondary forest succession require dif- ferent amounts of time, the changes in species diversity that ‘occur in each appear remarkably similar. Primary Succession at Glacier Bay We have already reviewed the basic patterns of primary suc- cession around Glacier Bay. Now we returnto Glacier Bay {0 examine successional changes in species diversity and com- position. William Reiners, lan Worley, and Donald Lawrence (1971) studied changes in plant diversity during succession at Glacier Bay. They worked at sites carefully chosen for simi- Jarity in physical features but differing substantially in age. ‘Their eight study sites were below 100 m elevation, were on glacial tll, an unstratified and unsorted material deposited by a glacier, and all had moderate slopes. The study sites ranged in age. that is. time since glacial retreat, from 10 to 1,500 years. Their youngest site, which was approximately 10 years old, supported a pioneer community of scattered Epilabium, Equisetum, and Salix. Site 2 was about 23 years old and sup- ported a mix of pioneer species and clumps of Populus and, Dryas. Site 3, which was approximately 33 years old, sup- ported a mat of Dryas enclosing clumps of Salix. Populus, and Alnus. Site 4 was 44 years old and was dominated by a mat of Dryas with few open patches. Site 5. which was approximately 108 years old, was dominated by a thicket of Alnus and Safix with enough emergent Populus and Picea to form a partial canopy. Site 6 was a 20(-year-old forest of Picea. Using geological methods, Reiners and his col- leagues dated site 7 at 500 years and site 8 at 1,500 yeurs. Both sites were located on Pleasant Island, which because it is located outside the mouth of Glacier Bay had escaped the most recent glaciation, which had destroyed the forests