ISSN 1188-603X

No. 453 April 19, 2012 Victoria, B.C.
Dr. A. Ceska, P.O.Box 8546, Victoria, B.C. Canada V8W 3S2


From: David Giblin e-mail

This year's Botany Washington is being co-sponsored by the Washington Native Plant Society and the University of Washington Herbarium at the Burke Museum. Our goal for this year is to provide an outstanding opportunity for participants of all levels of botanical skill to explore the rich diversity of Central Washington's flora across a range of elevations through three tracks of field study: one designed for technical study of selected taxa, one for improving plant keying skills using the Flora of the Pacific Northwest, and the third designed for individuals with little or no botanical background.

Taxonomic Study

Participants in this track will be studying Castilleja on Colockum Pass with Mark Egger on Saturday and Eriogonum on Table Mountain with Pam Camp on Sunday. It will be assumed that participants in the taxonomic track will have botanical and/or professional backgrounds and wish to improve their knowledge and understanding of specific taxonomic groups with botanical experts on these groups.

Keying with Hitchcock

You will join others who wish to hone their technical keying skills using the Flora of the Pacific Northwest in the company of an accomplished botanist who will provide expert guidance as you identify species unknown to you by working through Hitchcock's technical keys. It will be assumed that you have basic knowledge of botanical terminology and basic keying skills utilizing the Flora of the Pacific Northwest or another comparable technical flora. Opportunities will be provided for group keying and individual keying. This will be field study. Site locations to be determined.

Central Washington Wildflowers

Participants in this track are interested in exploring the native flora of different regions in Washington in a more informal setting. Your wildflower study will include field trips on Saturday and Sunday to areas of local floral interest with botanists knowledgeable about the local flora.

Base Camp

The Lazy F Camp is situated on 110 acres of mixed forest in the Manastash Canyon of the eastern Cascade foothills just 20 minutes off I-90 and not far from Ellensburg. Accommodations are rustic and will be dormitory style. We have reserved three cabins and a lodge that will accommodate up to 40 individuals. Participants will bring their own sleeping bags and other weekend essentials for a camp experience. Six meals including Friday and Saturday evenings dinner, Saturday and Sunday breakfasts, and Saturday and Sunday sack lunches will be provided and are included as part of the Botany Washington package.

Evening programs

On Friday evening Dr. Richard Olmstead, University of Washington, will discuss how and why contemporary systematics research has impacted many families that are found in our flora. On Saturday evening we will learn about Indian paintbrushes from Mark Egger, who conducts research on Castilleja and is the author of the treatment of this genus for an upcoming volume in the Flora of North America series.

Why register?

Botany Washington will provide botanists, ecologists, conservation biologists and other professionals with access to experts and an opportunity for in-depth study of selected taxonomic groups. It is also an opportunity for individuals new to the Flora of the Pacific Northwest to gain additional practice in technical keying with the support of expert guidance. The interested enthusiast wishing to learn more about regional floras of Washington will have two days in the field with knowledgeable local experts. Finally, it is an opportunity for botanists and others interested in our native flora to work together in a shared community of botanical learning. Membership in the Washington Native Plant Society is not required.


The cost of the weekend will be $225. This includes two nights rustic lodging, six meals and the opportunity to explore the flora of Kittitas County with knowledgeable botanists who will help you enhance your botanical skills no matter where on the botanical spectrum you reside.

Register online -

Mail-in registration form (contact David Giblin:

General questions should be referred to Catherine Hovanic or contact WNPS office at 206-527-3210, 1-888-288-8022 (toll free in Washington).


From: Roger del Moral

8:32 A. M. on Sunday, May 18, 1980 is a moment that is certainly etched in my mind and also in the minds of many others living in the Pacific Northwest and, if fact, around the world. At that moment, triggered by an earthquake, the fragile, bulging north side of this volcano collapsed. Within minutes, the landscape surrounding the mountain changed drastically. The north face became the largest recorded landslide and ultimately formed a flow of debris that overtopped nearby Mt. Margaret. This mass displaced Spirit Lake and then flowed west, filling the North Fork of the Toutle River valley. Almost simultaneously, the bottled up forces within the cone were released unleashing a massive lateral eruption that removed most life near the cone and toppled forests many kilometers distant and scorched trees even farther away. The cone had lost 400 m elevation and became a broken amphitheater.

Several intensely hot, glowing clouds of gas called pyroclastic flows roiled directly from the throat of the volcano thus sterilizing the land. As the day wore on, a cloud of pumice ascended 20 km and began to drift to the northeast. Near the cone, coarse pumice rocks fell to the new surfaces.

The texture of deposits became increasingly tiny with distance so that a fine dust called ash coated the landscape for several hundred km. The intense heat also caused glaciers and snowfields on the remaining sides of the crater to melt quickly. These caused great mudflows (called lahars) in the valleys rivers and creeks. The South Fork Toutle River lahar joined the debris flow of the North Fork and the combined mass reached the Columbia River. Deposits filled the river valley and several lakes were formed as creeks were dammed. The Muddy River lahar wreaked havoc on the southeast side of the cone, with deposits flowing into the Swift Reservoir to the south.

Together, these complex volcanic events set into motion an ecological response that has been the focus of research by many groups over the decades. Since that day, I redirected my research to focus on how plants could reclaim new and drastically altered landscapes. The research that I led with the invaluable assistance of my students and colleagues has changed how primary succession is understood and has had significant impact on restoration ecology.

Between 1980 and 2010, over 31 growing seasons, I studied the patterns and mechanisms of vegetation recovery in many habitats. The most interesting are those where no plants survived, so that recovery was, of necessity, by primary succession. Permanent plots and grids of contiguous plots formed the backbone of these studies, but along with many colleagues and students, I also conducted several broad surveys of vegetation across the northern flank of the cone (Pumice Plains) and the Muddy River. Several other studies involved local patterns of distribution and how species assembled into communities.

Here are some of the principal lessons learned:

1. Pioneer species were those that got there, not those that were best adapted to stress. Transplant studies showed that species establishing later in succession could survive and grow better than pioneers grow.

2. Initial establishment was improved by habitat amelioration by import of nutrients (e.g. dust, ballooning spiders), then by the development of "safe-sites" (e.g. rills and rock cracks). Over time, safe-sites degenerated as general conditions improved and competitors precluded establishment.

3. Species of relict and undisturbed sites adjacent to new surfaces do not materially hasten succession. Distances of only a few meters significantly reduce the rates of colonization. Further, species of such refuges are not suited to the newly created stressful conditions. Thus, both species richness and total vegetation cover decrease markedly over short distances.

4. Species accumulated at a much faster rate than did vegetation cover. Species interactions on primary surfaces were therefore slow in developing. Colonization was typically by a few successful individuals that quickly produced seeds on site, leading to their expansion.

5. This initial colonization was largely stochastic and results in heterogeneous vegetation such that adjacent plots often have low similarity. There were strong spatial auto-correlation effects when there were no dispersal barriers between closely located plots. Over time, similarity increases, but initial effects often persist.

6. Persistent initial effects appear to lead to alternative community types occupying similar habitats. It is premature to determine if these alternatives will persist.

7. The rate of succession, measured by how quickly a site reaches equilibrium, is controlled by environmental stress; species turnover occurs as the vegetaŦtion becomes denser and dominance hierarchies develop only when large cover develops. In similar habitats (e.g. a mudflow), the rate of succession is retarded with elevation.

8. The trajectories (the sequence of species composition in a single site) of succession are not fully predicable, and a given state in 2010 may have resulted from several alternative pathways.

9. Vegetation in essentially similar habitats may be quite distinct initially (due to stochastic immigration) and converge to similar vegetation; however this pattern was not generally observed and processes that can lead to convergent trajectories (competition, tight coupling between species and habitat variables) remain weak in most cases.

10. Trajectories at higher elevations have not recapitulated those of lower ones, showing that environmental severity affects both the rate and direction of trajecŦtories. This observation suggests that common space-for-time substitution methods used in toposequence studies may be more suspect than commonly believed.

11. Vegetation patterns were poorly predictable from environmental factors in at least six separate studies conducted at scales ranging the landscape to small plots. Predictive factors were more likely to be geographic, suggesting the importance of dispersal effects.

12. In a study of local depressions conducted from 1993 to 2008, the stochastic nature of community assembly was demonstrated. Each depression collected an essentially random input of species and collections of species were random in the early years. Starting with 2002, Lupinus lepidus expansion in many plots led to facilitation and inhibition, the development of consistent dominance hierarchies in many samples and the development of more homogeneous vegetation. There was a shift in statistical correlations between vegetation and predictor variables from purely spatial ones to habitat factors including soil moisture and pH. The overall relationship increased to 36%, a modest value in such studies. The effects of lupines vary between years because this species is subject to dramatic population fluctuations.

13. Landscape effects are extremely important. The nature of succession depends on the habitat, but also to a large extent on the availability of colonists. The importance of availability has been widely underappreciated.

14. Many of these findings are being applied in recovery studies of other volcanoes and in restoration of severely impacted ecosystems.


Carey, S.A., J. Harte & R. del Moral. 2006.
Effect of community assembly and primary succession on the species-area relationship in disturbed systems. Ecography 29:866-872.
Carey, S.A., A. Ostling, J. Harte & R. del Moral. 2007.
Impact of curve construction and community dynamics on the species-time relationship. Ecology 88: 2145-2153.
del Moral, R. 1981.
Life returns to Mount St. Helens. Natural History 90(5): 36-49.
del Moral, R. 1983.
Initial recovery of subalpine vegetation on Mount St. Helens, Washington. American Midland Naturalist 109: 72-80.
del Moral, R. 1993.
Mechanisms of early succession on Mount St. Helens. P. 79-100, in, J. Milne and D. W. H. Walton (eds.) Primary Succession on Land. Blackwell, London. [Summary paper]
del Moral, R. 1998.
Early succession on lahars spawned by Mount St. Helens. American Journal of Botany 85: 820-828.
del Moral, R. 1999a.
Predictability of primary successional wetlands on pumice, Mount St. Helens. Madroņo 46: 177-186.
del Moral, R. 1999b.
Plant succession on pumice at Mount St. Helens. American Midland Naturalist 141: 101-114.
del Moral, R. 2000a.
Succession and species turnover on Mount St. Helens, Washington. Acta Phytogeographica Suecica 85: 53-62.
del Moral, R. 2004.
How Lupinus Lepidus affects primary succession on Mount St. Helens. Pp. 208-215 in: E. van Santen (ed.) Wild and Cultivated Lupins from the Tropics to the Poles. Proc. 10th International Lupin Conference, Laugarvatn, Iceland, 2002. International Lupin Association, Canterbury, New Zealand.
del Moral, R. 2007.
Vegetation dynamics in space and time: an example from Mount St. Helens. Journal of Vegetation Science (Invited paper) 18: 479-488.
del Moral, R. 2009.
Primary succession on Mount St. Helens, with reference to Surtsey. Surtsey Research 12: 151-155. (Invited paper)
del Moral, R. 2009.
Increasing deterministic control of primary succession on Mount St. Helens, Washington. Journal of Vegetation Science 20: 1145-1154.
del Moral, R. 2010.
The importance of long-term studies of recovery after the eruption of Kasatochi Island. Arctic, Antarctic and Alpine Research 42:335-341.
del Moral, R. 2010.
Thirty years of permanent vegetation plots, Mount St. Helens, Washington (Data Paper). Ecology 91:2185.
del Moral, R. & L.C. Bliss. 1993.
Mechanism of primary succession: Insights resulting from the eruption of Mount St. Helens. Advances in Ecological Research 24: 1-66. [Summary Paper]
del Moral, R. & C.A. Clampitt. 1985.
Growth of native plant species on recent volcanic substrates from Mount St. Helens. American Midland Naturalist 114: 374-383.
del Moral, R. & S.Y. Grishin. 1999.
The consequences of volcanic eruptions. Chapter 5 in L.R. Walker (ed.) Ecosystems of Disturbed Ground, Ecosystems of the World Series (D.W. Goodall (Editor-in-Chief), Elsevier Science, Amsterdam.
del Moral, R & C.C. Jones. 2002.
Early spatial development of vegetation on pumice at Mount St. Helens. Plant Ecology 162: 9-22.
del Moral, R., J.E. Sandler & C.P. Muerdter. 2009.
Spatial factors affect primary succession on the Muddy River Lahar, Mount St. Helens, Washington. Plant Ecology 202: 177-190.
del Moral, R., J.M. Saura & J.N. Emenegger. 2010.
Primary succession trajectories on a barren plain, Mount St. Helens, Washington. Journal of Vegetation Science 21:857-867, plus supplements. [Key paper]
del Moral, R. & A.J. Eckert. 2005.
Colonization of volcanic deserts from productive patches. American Journal of Botany 92: 27-36. [Key paper]
del Moral, R. & E.E. Ellis. 2004.
Gradients in heterogeneity and structure on lahars, Mount St. Helens, Washington, USA. Plant Ecology 175: 273-286. [Key paper]
del Moral, R. & I.L. Lacher. 2005.
Vegetation patterns 25 years after the eruption of Mount St. Helens, Washington. American Journal of Botany 92:1948-1956.
del Moral, R. & L. Rozzell. 2005.
Effects of lupines on community structure and species association. Plant Ecology 180: 203-215. [Key paper]
del Moral, R., L.A. Thomason, A.C. Wenke, N. Lozanoff & M.D. Abata. 2012.
Primary succession trajectories on pumice at Mount St. Helens, Washington. Journal of Vegetation Science 23: 73-85. [Key paper]
del Moral, R., J.H. Titus & A.M. Cook. 1995.
Early primary succession on Mount St. Helens, USA. Journal of Vegetation Science 6: 107-120.
del Moral, R. & L.R. Walker. 2007.
Environmental Disasters, Natural Recovery and Human Response. Cambridge: Cambridge University Press.
del Moral, R., L.R. Walker & J.P. Bakker. 2007.
Insights gained from succession for the restoration of structure and function. Chapter 2 in: L.R. Walker, J. Walker & R.H. Hobbs, Linking Restoration and Succession in Theory and in Practice. New York: Springer.
del Moral, R., D.M. Wood & J.H. Titus. 2005.
How landscape factors affect recovery of vegetation on barren surfaces. Pp. 93-10, in Mount St. Helens 20 years after recovery (V.H. Dale, F. Swanson & C. Crisafulli, eds.). [Summary paper]
del Moral, R. & D.M. Wood. 1988a.
Dynamics of herbaceous vegetation recovery on Mount St. Helens, Washington, USA, after a volcanic eruption. Vegetatio 47: 11-27.
del Moral, R. & D.M. Wood. 1988b.
The high elevation flora of Mount St. Helens, Washington. Madroņo 35: 309-319.
del Moral, R. & D.M. Wood. 1993a.
Understanding dynamics of early succession on Mount St. Helens. Journal of Vegetation Science 4: 223-234. [Summary paper]
del Moral, R. & D.M. Wood. 1993b.
Early primary succession on a barren volcanic plain at Mount St. Helens, Washington. American Journal of Botany 80: 981-991.
Dlugosch, K. & R. del Moral. 1999.
Vegetational heterogeneity along environmental gradients. Northwest Science 43: 12-18.
Fuller, R.N. & R. del Moral. 2003.
The role of refugia and dispersal in primary succession on Mount St. Helens, Washington. Journal of Vegetation Science 14: 637-644.
Grishin, S.Y., R. del Moral, P.V. Krestov & V.P. Verkholat. 1996.
Succession following the catastrophic eruption of Ksudach volcano (Kamchatka, 1907). Vegetatio 127: 129-153.
Grishin, S.Y., & R. del Moral. 1996.
Devastation and restoration of subalpine Betula ermannii forest after the 1975 Tolbachik eruption, Kamchatka Peninsula. Botanichesky Zhurnal (Russian).
Marler, T.E. & R. del Moral. 2011.
Primary succession along an elevation gradient 15 years after the eruption of Mount Pinatubo, Luzon, Philippines. Pacific Science 65(2): 157-173.
Titus, J.H. & R. del Moral. 1998a.
Vesicular-arbuscular mycorrhizae influence Mount St. Helens pioneer species in greenhouse experiments. Oikos 81: 495-510.
Titus, J.H. & R. del Moral. 1998b.
Seedling establishment in different microsites on Mount St. Helens, Washington, USA. Plant Ecology 134: 13-26.
Titus, J.H. & R. del Moral. 1998c.
The role of mycorrhizae in primary succession on Mount St. Helens. American Journal of Botany 85: 370-375.
Titus, J.H., R. del Moral & S. Gamiet. 1998.
The distribution of vesicular-arbuscular mycorrhizae on Mount St. Helens, Washington. Madroņo 45: 162-170.
Titus, J.H., P. J. Titus & R. del Moral. 1999.
Wetland development in primary and secondary successional substrates fourteen years after the eruption of Mount St. Helens, Washington, USA. Northwest Science 73: 186-204.
Tu, Mandy, J.H. Titus, R. del Moral & S. Tsuyuzaki. 1998.
Composition and dynamics of wetland seed banks on Mount St. Helens, Washington, USA. Folia Geobotanica 33: 3-16.
Tsuyuzaki, S. & R. del Moral. 1995.
Species attributes in early primary succession on volcanoes. Journal of Vegetation Science 6: 517-522.
Tsuyuzaki, S., J.H. Titus & R. del Moral. 1997.
Seedling establishment patterns on the Pumice Plain, Mount St. Helens, Washington. Journal of Vegetation Science 8: 727-734.
Walker, L.R. & R. del Moral. 2003.
Primary succession and Ecosystem Rehabilitation. Cambridge University Press, Cambridge, UK.
Wood, D.M. & R. del Moral. 1987.
Mechanisms of early primary succession in subalpine habitats on Mount St. Helens. Ecology 68:780-790. [Key paper]
Wood, D.M. & R. del Moral. 1988.
Colonizing plants on the Pumice Plains, Mount St. Helens, Washington. American Journal of Botany 75:1228-1237.
Wood, D.M. & R. del Moral. 2000.
Seed rain during early primary succession on Mount St. Helens, Washington. Madroņo 47: 1-9.

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