April 1986 FREMONTIA A journal of the California Native Plant Society -r: FREMONTIA Vol. 14 No. 1 April 1986 Copyright © 1986 California Native Plant Society Phyllis M. Faber, Editor Laurence J. Hyman, Art Director Beth Hansen, Designer MATERIALS FOR PUBLICATION Members and others arc invited to submit material for publication in Fremontia and the Bulletin. All time-value material should be addressed to the Bulletin. Fremontia is a journal for laymen about California plants. Technical botanical articles should be directed to other more scholarly journals. Please double-space copy, using wide margins and fresh typewriter ribbon, on 8'/2-by-ll paper, and include name, address, and phone number, and submit two copies of ms. As a general rule, in the interest of consistency, botanical nomenclature will conform to Munz, A California Flora. Please identify each plant referred to by its botanical name and, if there is one, by its common name. Photographs should be black-and-white glossy prints, prefer- ably 8-by-lO size or accompanied by negatives. Authors are urged to submit MS on floppy discs, produced via Wordstar or Multimate soft- ware on an IBM compatible computer. THE COVER: A mountain lake southwest of Rush Creek in the Sierra Nevada, photographed by Victor Duran in July 1924. Duran's photographs arc featured in this issue of Fremontia. MEMBERSHIP Dues include subscriptions to Fremontia and the Bulletin. Life (Individual/Couple. $450/500 Individual or Library. $18 Supporting................. $50 Student or Retired. .. $12 Household................. $30 Retired Couple......$15 ADDRESSES Memberships; Address Changes; Officers; General Society Inquiries; Conservation Trust Fund: CNPS, 909 12th St., Suite 116, Sacramento, CA 95814. (916) 447-CNPS Fremontia (Editor): Phyllis M. Faber, Editor, 212 Del Casa Drive, Mill Valley, CA 94941. (415) 388-6002 Fremontia (Advertising): Nancy Dale, Rancho Santa Paula #7,500 W. Santa Maria, Santa Paula, CA 93060. (805) 525-6319. Bulletin: Pauleen Broyles, Editor, P.O. Box 763, Paradise, CA 95969-0763 CNPS Botanist, Data Base: Rick York, 909 12th St., Suite 116, Sacra- mento, CA 95814. (916) 324-3816 or (916) 447-CNPS EXECUTIVE COUNCIL President................................Charlice Danielsen Vice President, Administration................Laurie Kiguchi Vice President, Finance.....................R. Arthur Hayler Vice President, Conservation.......................Ken Berg Vice President, Legislation........................Bob Berka Vice President, Rare Plants................James P. Smith, Jr. Vice President, Publications....................Harlan Kessel Legal Advisor................................Scott Fleming Recorder..................................Joanne Kerbavaz Corresponding Secretary.....................Susan Sommers Past President..................................Robert Will DIRECTORS-AT-LARGE Richard Burgess, Jim Dice, Roman Gankin, Jo Kitz, Mary Merryman, Tim Thomas California Native Plant Society Dedicated to the Preservation of the California Native Flora The California Native Plant Society is an organization of laymen and professionals united by an interest in the plants of California. It is open to all. Its principal aims are to preserve the native flora and to add to the knowledge of members and the public at large. It seeks to accomplish the former goal in a number of ways; by monitoring rare and endangered plants throughout the State; by acting to save endan- gered areas through publicity, persuasion, and, on occasion, legal action; by providing expert testimony to governmental bodies; and by supporting financially and otherwise the establishment of native plant preserves. Much of this work is done through CNPS Chapters throughout the State. The Society's educational work includes: pub- lication of a quarterly journal, Fremontia, and a quarterly Bulletin which gives news and announcements of Society events and conser- vation issues. Chapters hold meetings, field trips, and plant and poster sales. Non-members are welcome to attend. The work of the Society is done by volunteers. Money is provided by the dues of members and by funds raised by chapter plant and poster sales. Additional donations, bequests, and memorial gifts from friends of the Society can assist greatly in carrying forward the work of the Society. Dues and donations are tax-deductible. CHAPTER PRESIDENTS (AND DIRECTORS) Bristlecone (Inyo-Mono)....................Doris Fredenhall Channel Islands..............................Darl Dumont Dorothy King Young (Gualala)..............Robert Z. Perkins Kern County................................Diane Mitchell Marin.......................................Sue Hossfeld Milo Baker (Sonoma County)....................Liz Parsens Monterey Bay..................................Vern Yadon Mount Lassen.................................Mike Foster Napa..........................................Joe Callizo North Coast.....................................Ken Berg Northern San Joaquin Valley (Modesto)..........John Herrick Orange County...............................Dave Bramlet Riverside/San Bernardino Counties.............Andy Sanders Sacramento Valley......................Lorraine Van Kekerix San Diego..................................Andrew Wilson San Francisco Bay.........................William R. Keeler San Gabriel.................................Harry Spilman Sanhedrin (Ukiah).............................Mark Albert San Luis Obispo............................Gary Ruggerone Santa Clara Valley.............................Bart O'Brien Santa Cruz...............................Adrienne Harrold Santa Monica Mountains.................Linda Hardie-Scott Sequoia (Fresno).............................Jeanne Larson Shasta........................................Jim Mallory South Coast (Palos Verdes)...................Richard Dulzeul Tahoe.........................................Bob Allard 2 rrfSH^HSW.. ¦¦•;**>'¦ /tiSStev A bay at the north end of Cedros Island. Photographs by the author. BAJA CALIFORNIA'S PACIFIC ISLAND JEWELS by Thomas Oberbauer Some of California's most fascinating floristic areas are isolated mountains, coastal closed-cone pine forests, and islands. Two locations off the coast of central Baja California possess all of these attributes: Guadalupe and Cedros islands. Guadalupe Island and Cedros Island are similar in many respects. They are roughly comparable in size: Guadalupe Island is twenty-two miles long with a total of ninety-eight square miles; Cedros Island is twenty miles long with a total of 134 square miles. Both have mountain peaks approaching 4,000 feet elevation, and because of the regional wind patterns, both receive con- ;derable moisture from fog on their otherwise dry slopes. Guadalupe and Cedros islands have another similarity. They are part of the California floristic region. Both are farther south than the mainland occur- rence of this floristic region; however, they share a number of species with California's Channel Islands and the mainland of California and northern Baja Cal- ifornia. Cedros is clearly a desert island; its lower slopes receive only about two and one-half inches of rain each year, and the vegetation is an open, brushy and spiny community of cacti, agaves, and low shrubs, yet the high points support chaparral and coastal sage scrub and a picturesque pine forest. The pine forest grows on the 3 north end of the island, precisely where spring and summer fogs prevail The vegetation on Cedros, even in the dry southern areas, is amazingly diverse. Desert canyons contain such unusual-looking shrubs as the large, silver-leaved sunflower, Viguiera lanata, elephant trees or torote (Pachycormus discolor), and a variety of less striking shrubs, including the bursage {Ambrosia chenopodiifolia) and the small-flowered, delicate Cedros sage (Salvia cedrosensis). Guadalupe Island also supports a sparse, xeric vege- tation on its lower slopes. It receives an average of five inches of rain each year. The upper slopes toward the north end of the island also support a pine forest in addition to cypress and palm woodlands. Unfortu- nately, Guadalupe Island has suffered the ravages of feral goats, and it is difficult in many places to identify the original vegetation because the soil surface has been denuded. In other areas, particularly the northern end and at higher elevations where feral goats have concen- trated, introduced species have become widespread. Huge stands of the South American tree tobacco (Nico- tiana glauca) and carpets of weedy grass, wild oats (Avena fatua), and wild barley (Hordeum leporinurri) cover large areas. Cedros, on the other hand, appears little affected by feral browsers and grazers. There the vegetation is apparently adapted to the browsing of a few endemic mule deer. There are other significant differences between Guadalupe and Cedros islands. Cedros Island is a geo- logic mix of rock types, including sedimentary forma- Canyon vegetation on Cedros Island is more abundant than on the dry, exposed slopes which receive only about two and one-half inches of rain a year. tions, metamorphosed shales, volcanics, granitic rock, and even some serpentine. As such, it is a piece of the continental borderland that has become isolated through time and undoubtedly was connected to the mainland peninsula in the geologic past. It is now only fourteen miles from the mainland and isolated by shal- low water. In contrast, Guadalupe Island is composed purely of volcanic rock outside the edge of the continen- tal borderland and is surrounded by water two and one- third miles deep. It lies 150 miles from the mainland and was never connected to it, existing as a truly oceanic island. Both Cedros and Guadalupe are scenically impres- sive, with high cliffs, steep slopes, and deep canyons. Under pristine conditions, Guadalupe Island must have been fantastic, but it now remains as a memorial to the destructive nature of feral goats. Cedros Island has some heavily disturbed areas, yet it still supports an array of species and a unique assemblage of communities. Endemic and Disjunct Species Both islands are rich in endemic plant forms, and some of the most notable are the pines. Both support two needle pines that, depending on taxonomic opin- ion, are either varieties of Bishop pine (Pinus muricata, Santa Cruz Island pine (P. remorata), or Monterey pine (P radiata var. binata on Guadalupe and P. radiata var. cedrosensis on Cedros). These pines are related to pines on the northern Channel Islands, central California, and one location in northern Baja California. The pines on Cedros Island grow in striking contrast to the surrounding desert vegetation in three main popu- lations: one isolated stand in the middle and two stands separated by a low saddle on the northern end of the island. These forests appear to be healthy and vigorous in an atmosphere that is heavy and laden with moisture, even when free of fog and clouds. The pine forest on Guadalupe Island is dying. It con- sists of scattered large, old trees, predominantly on cliffs, mixed with decaying old trunks and remnant skeletons. Guadalupe Island's forest evidently was more developed, but goat predation has eliminated seedling production of the pines as well as the endemic palm, Brahea edulis, and the endemic cypress, Cupressus guadalupensis. Cedros Island contains fifteen species and varieties of plants that apparently are endemic even though it lies only a short distance from the mainland. Its montane nature accounts for some that do not have similar habitats on the mainland. However, several of Cedros Island's endemics grow in habitats at low elevations. These might be expected to grow also on the adjacent mainland, but they have not yet been discovered there. Some of the most interesting include the wild fuchsia {Xylonagra arborea van arborea), a prickly phlox (Lep- todactylon veatchii), Harfordia fruticosa, which is a polygonaceous shrub with red lantern-shaped pods, and several endemic dudleyas, especially the recently described stump-leaved Dudley a pachyphyta. Guadalupe Island supports the largest number of endemics (thirty-five taxa) of any California or Baja California island. Several are now thought to be extinct, including a tree in the olive family (Hesperalaea) and two mints, Pogogyne tenuiflora and Saturejapalmeri. Several other species have been found only within the crater of an islet near the southern end of Guadalupe Island. Additional endemic species probably occurred on Guadalupe Island; however, they have since become extinct because of the introduction of goats. Many of Guadalupe Island's endemics are showy and would make fine ornamentals in frost-free gardens, including the pink-flowered, succulent-leaved Talinum guadalupense, a yellow-flowered shrub reminiscent of dusty miller, Perityle incana, and a yellow-flowered amaryllidaceous plant, Triteleia guadalupensis. The endemic palm, Brahea edulis, has already shown up in a few specialized gardens. Both Cedros and Guadalupe islands share a number of species with California's Channel Islands and the mainland far to the north. For example, Guadalupe Island supports the giant coreopsis (Coreopsis gigantea) in common with the northern Channel Islands and coastal Santa Barbara County; Camissonia guadalupen- sis in common with San Clemente Island (though a dif- ferent variety); tree-mallow (Lavatera occidentalism with the Coronado Islands near Rosarita Beach; and Esch- scholzia ramosa, the island poppy, with all of the other Channel Islands, including Cedros. In total, Guadalupe Island shares twenty species of insular endemics with the other Channel Islands. Such plants as mission manzanita (Xylococcus bicolor), silk-tassel (Garrya veatchii), and chamise (Adenostoma fasciculatum), found on Cedros Island, also occur miles to the north on the mainland. Chamise and mission manzanita also occur on the California Channel Islands. Guadalupe Island supports, or supported, a few plants with unusual distributions, including species dis- junct to mainland central California several hundred miles north. Such species are a polypoid fern (Polypo- dium scouleri), woolly yarrow (Eriophyllum lanatum var. grandiflorum), and a red-flowered currant (Ribes sanguineum). It is conceivable that these species existed on the mainland south of their modern distribution and closer to Guadalupe Island during past geologic times. How- ever, even if they occurred on the adjacent mainland, they would have had to disperse across 150 miles of ocean to reach the island. The disjunct species appear able to persist on Guada- lupe and Cedros islands because of the supplemental moisture they receive from fog and a cool marine envi- ronment. The topographic diversity of the islands is also an important factor, since the fog drip would not be as prevalent on islands with lower elevations. Natural Laboratories Guadalupe and Cedros islands are key indicators of past climatic conditions and dispersal mechanisms for plants. Their vegetation and plant communities raise a number of questions: How did the pines and disjunct species find their way there? And what about the endemics? Answers to such questions are continually being refined. There were times, when conditions were more favorable, when pines similar to those on Cedros and Guadalupe islands existed much further down the coast than they do at present. They may then have dis- persed overland in the case of Cedros Island and over water in the case of Guadalupe Island, becoming estab- lished at lower elevations when rainfall was greater. The isolated populations in the fog drip zones are remnants of former populations that retreated from sea level as the climate became drier. Similar dispersal may have taken place from the north for other plants with disjunct distributions. The endemics may have either evolved on the islands or dispersed to them from the mainland and persisted after the mainland populations died out. Recently evolved species would be expected to be closely related to mainland forms, such as the endemic Cryptantha foliosa and the lupines on Guadalupe Island, while older species that persisted on the islands but died out on the mainland may be unrelated to mainland species today. An example of the latter would be the extinct Hesper- alaea, Galvesia speciosa, and Crossossoma on Guada- lupe Island. Crossossoma, long believed to be an insu- lar endemic, was recently discovered in the Santa Monica Mountains. Its distribution is similar to Cata- lina ironwood (JLyonothamnus spp.) from the Channel Islands, which has been found in mainland fossils and must have been more widespread on the mainland at one time. Both of these islands have strong ties to Alta Califor- nia and the California Channel Islands. They are rich in natural habitats that raise some botanical questions and provide answers to others. Much of the botanical value of these islands will be eliminated unless the goats are removed from Guadalupe Island and constant efforts are made to prevent introduction of plant-eating mammals on Cedros Island. It would be worthwhile to educate people on both sides of the border about the importance and sensitivity of these island jewels. 5 GENE CONSERVATION IN CALIFORNIA'S FORESTS by Constance I. Millar The University of California's Wildland Resources Center has established a new program of forest gene conservation to ensure that California's rich and diverse forests maintain their vigor and productivity in the face of human activities. At an international level, conser- vation biologists recognize the importance not only of protecting rare species from extinction by safeguarding habitats, but of caring for the genetic diversity of estab- lished forests. In some parts of the world past neglect, insufficient knowledge of genetics, and indiscriminate forestry practices have resulted in forests that are impoverished by a loss of genetic diversity. These prac- tices include repeated harvesting of the best trees, leav- ing poorer specimens to reproduce; planting species and individuals that are poorly adapted to local soil and cli- matic conditions; allowing exotic weed trees to escape into forests; and excessively logging areas with steep slopes or unusual soils. All of these practices result in reduced levels of productivity and lowered long-term genetic adaptability, including greater susceptibility to infestations of pests. Recognizing this vulnerability of trees to man's interventions, forest geneticists have mounted a worldwide effort to protect threatened and sensitive forest areas and to prevent future misuse of forests still in wild condition. In California we have benefited from knowledge gained in forests around the world. We are also for- tunate in that many California forests retain the genetic diversity expected under natural conditions. Neverthe- less, many forested areas in California are threatened by forestry practices that will erode their genetic quality. Professional foresters and forest geneticists from pri- vate industry, the California Department of Forestry, and the USDA Forest Service's Tree Improvement Cen- ter and Institute of Forest Genetics are working together to protect the genetic diversity of our forests. Some of the techniques they use include reforesting areas, where possible, with locally collected seed; establishing breed- ing zones so that seeds are collected for a specific area only from areas with closely related elevations; and pre- serving an assemblage of species resembling that found under natural conditions. These groups have integrated guidelines for gene conservation into their forest man- agement strategies and sponsor gene conservation research. At least two private groups, the National Council on Gene Resources, a research and educational organiza- tion, and GenRec, a consulting firm, specialize in gene conservation analysis. The California Native Plant Soci- ety is dedicated to gene conservation through the pro- tection of rare and endangered species and habitats. Currently, gene conservation programs follow two Sequoiadendron giganteum (giant sequoia) growing in King's Canyon National Park provides an example of a forest tree to be protected through the University of California gene conservation program. main approaches: ex situ and in situ management. The former usually consists of collecting and storing seeds and pollen in long-term storage banks or of creating tree plantations by planting genetically identical specimens cloned by tissue culture or by cuttings. These ap- proaches preserve genes in a static condition (as seeds or clones) and freeze natural evolutionary processes. In the future, if there is a need to return to the genetically diverse parent stock, the stored seeds or established clones can be grown into trees and the preserved genes integrated into ongoing forest management programs. The in situ approach recognizes both that species exist in interbreeding communities subject to natural selec- tion and that natural evolutionary processes tend to maintain genetic variability of individuals and species. Thus it emphasizes the value of preserving minimally managed areas where species are subject to natural pro- cesses, both physical and biological, and genetic diver- sity is more likely to be retained. A new program of forest gene conservation was begun in summer 1985 on the Berkeley campus of the University of California. The project, under the direc- tion of the Wildland Resources Center and financed by the California Environmental Protection Program (the automobile license plate fund), will provide information and resources to protect the genetic diversity of Califor- nia's forest flora. The program consists of four projects, each of which will address a specific problem in gene conservation. The first will prepare a computer-based Pinus radiata (Monterey pine) growing on Monterey peninsula provides an example of a forest tree to be protected through the University of California gene conservation program. Hedges of Pinus radiata, used in ex situ cloning at the Russell Reservation, are clipped annually to produce genetically identical shoots that are easily rooted. catalog of lands in California where in situ forest gene conservation can occur. Although many acres in Cali- fornia are allocated for preservation in national forests and national and state parks, the extent of protection of forest species and communities is unknown. The catalog will provide information on the adequacy and types of protection provided by existing preserves and will identify gaps in protection of forest gene resources. The second project will restore valuable ex situ gene conservation collections that have suffered due to lack of funds for maintenance. Seed collections and clonal plantations of several important species, including Mon- terey pine, coast redwood, and giant sequoia, will be inventoried and renovated by new cloning and creation of new plantations. A long-term maintenance program will be established. The third project will complete seed collections in California for long-term storage from the ranges of Douglas fir and ponderosa pine. These important tim- ber species are among the most productive and valua- ble forest trees in the world and at present contain vast stores of genetic variation that should be protected for the future. An extensive seed collection also will be made for Brewer spruce, an endemic species limited to a few locations in northern California and southern Oregon. The fourth project of the forest gene conservation program will undertake extensive studies of geographic patterns of genetic variation in ponderosa pine, Douglas fir, and Brewer spruce. Seeds will be analyzed with a standard laboratory technique of electrophoresis, which shows protein diversity, used as an indicator of genetic diversity. Such information will allow wise decisions to be made on the nature and adequacy of a gene protec- tion program for these species. It is anticipated that the University of California's gene conservation program will continue for several years and that it will provide the tools for genetic con- servation of important tree species in California. 7 THE PHOTOGRAPHY OF VICTOR DURAN by Jacob Sigg Victor Duran, explorer, plant collector, and photog- rapher, was born in 1897 and spent much of his boyhood biking and collecting butterflies in the mountains around Los Angeles. After receiving a master's degree in physics from the University of California, he became a student of botany under the guidance of Dr. Willis Lynn Jepson. Duran pursued a study of the floristic composition of the White Mountains from 1926 to 1933, making numerous trips into remote areas of the moun- tains with horse or mule to collect plants. This collec- tion later formed the basis for the Flora of the White Mountainshy R.M. Lloyd and R.S. Mitchell (1973). Two plants, Heuchera duranii Bacigalupi and Streptanthus cordatus Nutt. var. duranii Jepson, were named in his honor. During this period Duran also made an extensive col- lection of photographs of the Sierra and the White Mountains, some on glass plates, others on the new cel- luloid film. This collection is intact today, together with meticulous field and photographic notes, and is admin- istered by his daughter, Sylvia Duran Charnoff. She shares her father's interest in photography, and in lichens and mushrooms, and has been a past contribu- tor to Fremontia. The photographs for this essay were Left: Snow on Boundary Peak, White Mountains, June 1932. Photographer's log for the same picture taken 800 feet lower reads: "Trail Creek, White Mountains, at about 8,200 feet. Pinyons on near slope; willow, birch, aspen in canyon; meadow and rabbit brush on flats; sagebrush on slopes. Boundary Peak." The pines in the distance occupy higher ground and would be either bristlecone or limber pine. Below: View up Merced Canyon from Cloud's Rest in Yosemite National Park, July 1922. Scraped clean by the upper trunk of the Merced glacier, small quantities of rock particles and humus collect and allow the establishment of forests of white fir, red fir, western white pine, and lodgepole pine. selected from the Duran collection and, to a degree, reflect the photographer's preoccupation with the coniferous trees of California. Victor Duran joined the Scientific Photography Laboratory at the University of California, Berkeley, in 1933 and in time became principal scientific photogra- pher, a position he held until his retirement a number of years ago. His contribution to research efforts and high-quality publications at the university and to the flora of the White Mountains are universally hailed by his many friends. Simpson Meadow at 6,000 feet with Mt. Woodworth in the background on the Middle Fork of Kings River, Sierra Nevada, 1926. Pines, even when young, require full exposure to the sun. In this meadow they would normally be outcompeted at the seedling stage by grasses and shrubs. Because they managed to survive that dangerous time, they are enjoying the extra water available in the meadow and are growing with unexcelled vigor. 9 Above: A large ponderosa pine on Dutch Flat near Idylwild in the San Jacinto Mountains with Carl Meyer standing next to it, May 1932. Acorn woodpeckers drill holes in the thick bark of some trees for storage of seeds. The holes drilled are exactly the same size as the seeds so they cannot be removed by marauding squirrels. In seed-short years woodpeckers have been known to pound pebbles into the holes instead. Above right: Sheep grazing around Jeffery pine at end of road below Morgan Creek near Round Valley, Inyo County, July 1932. Adapted to poor soils, cold and heat, and great diurnal variations in temperature, Jeffrey pine is the dominant tree above the pinyon zone on the eastern slopes of the Sierra. Winter winds blow large quantities of snow into the deep canyons, which drop off sharply from the Sierra crest. This snow-melt enables trees to attain large size where sufficient soil can be found. Right: Western juniper above Morgan Creek near Round Valley in Inyo County, July 1932. John Muir was not fond of the western juniper (Juniperus occidentalis var. australis), which he considered too somber. Its toughness in exposed, ungrateful conditions, its ability to wrest a living from rock, its great longevity, and its picturesque form inspire fascination and awe in the observer. This is an exceptionally tall and lush juniper, growing on a site on the east side of the Sierra that would normally be occupied by a more demanding species. 10 «fJ*/, $¦?*. Utah juniper at Cedar Flats, June 1932. Utah juniper (Juniperus osteosperma) is the near-constant companion of the pinyon pine throughout the high deserts of the Great Basin. The indicator plant for this area is the Great Basin sagebrush (Artemisia tridentata), shown here with a locoweed (Astragalus lentiginosus). 11 tf -'"i."'* '#'•*' *'/w 12 BK#3*,,"^v* 'Ur^.?'- *£'* i^S. ¦i'-J5!^5*. 1. / * I in-i ini ili«.- Inli;i'-'i'i'l linihci piik-mi liiuiiul;ii\ K'.ik in ilk' W Ink-Mnimi.iin^. I'"2 WiJc^iik'.nl in »i'»k'in Ni'iili \nu'ik'.i. [lie lllllK'1 pill. \l'lll!f. '/.'Wi'M III ( .lllllHlll.1 .Mill V'\:kl.l il.i-lipk'- uiil.. ilk' Inj'lk^i .k'V.ilions \il.ipk\l iniliv mill. II OMiipck'- Willi ilk' 1'IIMk'l.illk' liM ll.ll'll.ll. hill lll'l k'l lllll'Jl'\ll\ I llllKl ptlk'> Ik'U'l -1.0111 111 L'lllW i'lllk'1 \C1\ l.li'Jl i'I WIN I'M 13 Right: Boundary Peak in the White Mountains viewed from Mustang Peak, June 1932. The rockier, more angular peaks at the northern end of the White Mountain range lack the special conditions that obtain in the dolomitic soils at the southern end. The trees are much younger and less dramatic, and the proportion of limber pine to bristlecone pine increases. ^ Below: White Mountain peak showing between bristlecone pines on a ridge south of McCloud Camp, June 1926. The Great Basin bristlecone pine (Pinus aristata) finds a competitive advantage in the white dolomite soils of the White Mountains. Dolomite is unfavorable to the seedlings of sagebrush, which normally crowd out the pine seedlings. The oldest trees are usually found on this dolomitic limestone. r®Ci Right: Hill west of Emigrant Meadow in the Stanislaus National Forest, July 1924. Treeline at the northern end of the Sierra is lower than it is further south. Here the treeline occurs below glaciated areas. Note the volcanic rocks on right. Far right: A view of Montgomery Peak from a state highway near Queen, Nevada, on the California-Nevada line, June 1932. A treeless gap occurs between the upper range of the pinyon and the lower margin of bristlecone/limber pine belts, graphically illustrated here. 14 <- » ¦¦;¦ i*v-"i T* * h" f*-- *......^mm^fi^v- ¦>. -^ 15 16 ';?«JK1 * • ¦¦?-* \. "jIV ;¦»*-? # m V* Left: Victor Duran standing beside ponderosa pine one mile west of Pine Flats in the Sierra Madre Mountains, 1921. Of the one hundred species of pine, only the Scotch pine (Pinus sylvestris) has a greater geographic range than Pinus ponderosa. The range of the ponderosa extends throughout western North America. A confluence of favorable environmental factors enables it to reach its most noble dimensions in California. Below: A view within a Jeffrey pine forest at Sand Flat near Mono Mills, July 1932. A mix of young and middle-aged Jeffrey pines. On well-watered sites, such as these deep volcanic soils, the Jeffrey pine becomes very large. The clean, open, park-like stand of old trees is one of California's most beautiful forests. Opposite page, top: A pure stand of balsam root in a forest opening near Long Valley in the Warner Mountains, August 1936. Alluvial deposition creates a deep, sandy soil, beloved by balsam root (Balsamorhiza) and surrounded by aspen, fir, and pine. Opposite page, bottom: Magee Peak seen beyond a stand of hemlocks near Crater Peak, Lassen National Forest. Mountain hemlock (Tsuga mertensiand) is a conical tree whose branches have slender branchlets forming graceful pendant masses of foli- age. In youth and middle age it is charmingly feminine, becoming stately when older. It was John Muir's favorite Sierra tree. A peculiarity of the species is the mass of basal branches usually retained throughout the tree's life. 17 **V «*%*.** Digger pine growing east of the Pinnacles, April 1932. This vigorous Digger pine (Pinus sabiniana), sixty feet high and twelve feet in girth at four feet above the ground, is the only native tree (as opposed to shrubby pinyons) in the genus that forks into many branches instead of concentrating on a terminal leader. 18 A GRAND TOUR OF NORTHERN CALIFORNIA'S CONIFERS (Part I) by Glenn Keator Come on a tour of the northwestern corner of Cali- fornia, an outstanding place to see a rich diversity of conifer habitats and species. Some parts of this area are well known to the general public, but others are known only to more adventurous naturalists. California is renowned for its many fine conifers — coast redwood, giant sequoia, bristlecone pine, some fifty-eight taxa in all. Much of this richness results from a great diversity of topography, climate, and soils. And part of the story lies with the evolutionary history of species, many of which represent what botanists call relicts — plants that today are restricted to special environments because the climate has changed or they are poor competitors with more vigorous neighbors. Examples are found in nearly every genus of our coni- fers: Torrey pine (Pinus torreyana), several species of closed-cone pines, Santa Lucia fir {Abies bracteata), Brewer's spruce (Picea breweriana), California nutmeg An overview of the beach pine forest near Samoa. Photographs by the author. (Torreya californica), bigcone spruce (Pseudotsuga mac- rocarpa), and a number of cypresses (Cupressus spp.). We'll begin the tour with the better known coastal areas up around Eureka and Humboldt Bay in the heart of the best coast redwood country. A map of north- western California is all that is needed to follow our odyssey eastward from the luxuriant coastal forests, up the Smith River, across the varied and rugged mountains of the Klamath Range, and down the Sacramento River to the hot, dry interior of the Central Valley. Elevations range from sea level along the coast to around 7,800 feet at Little Duck Lake in the Klamath Range. Rocks and soils include granites, clays, silts, sands, volcanics, and serpentines. The area embraces more than one-half of all California's conifers, including the genera Pinus (pine: eleven species); Abies (fir: six species); Tsuga (hemlock: two species; Picea (spruce: three species); Pseudotsuga (Douglas-fir: one species); Taxus (yew: one species); Sequoia (redwood: one species); Juniperus (juniper: two species); Cupressus (cypress: one species); Chamaecyparis (false cypress: two species); and Calocedrus (incense-cedar: one species). To drive the full route, you should reserve several days, preferably a week or more, for the distances involved are several hundred miles. Some of the roads are well worn but usually passable for ordinary vehicles. The ideal time of year is between late June and late July when most of the snow has melted, roads are passable, and the profusion of interesting flowers is at a peak. Humboldt Bay Few tourists see more than the edge of Humboldt Bay as they pass through Eureka. A short side excursion along State Highway 255 takes you out to the sand spits around redwood milling plants near the hamlet of Samoa, where some of the best dune vegetation of Northern California contrasts strikingly with weathered beach pine forests growing on stabilized dunes. Beach pines (Pinus contorta) are easily recognizable by their rounded canopies, short, dark green needles in twos (with a slight twist in each needle), rugged bark, and lux- uriant crops of small, prickly, scaled seed cones. The seed cones vie with those of the closely related lodge- pole pine (P. murrayana) for being the smallest in the state, often no more than an inch or so long. P. mur- rayana is equally well adapted to wind-swept conditions and nutrient-poor, sandy soils but is found in the high mountains. 19 Under the pine canopy is a lush understory (owing to high rainfall and summer fogs) of kinnikinnick man- zanita {Arctostaphylos uva-ursi), ground lichens, rattle- snake orchid (Goodyera oblongifolia) with mottled leaves, and mosses, as well as evergreen shrubs such as wax myrtle (Myrica californica), silk-tassel bush (Garrya elliptica), salal {Gaultheria shallori), and California huckleberry (Vacciniwn ovatum). Many of the plants require cool, damp conditions and acid soils, conditions much like the northern boreal forests in which they probably evolved. To the north and west of Areata lie the nearly pristine Lanphere-Christensen Dunes, a preserve belonging to The Nature Conservancy. Permission to visit may be arranged through The Nature Conservancy, the preserve caretaker, or the biology department of Humboldt State University in Areata. This dune ecosystem presents the successional stages of vegetation associated with dune systems: from foredunes where the native dune ryegrass (Elymus mollis) helps hold down loose sand; across mature dunes with their rich array of flowers, including the bright yellow rare and endangered Menzies' wall- flower (Erysimum menziesii); to blowouts and hollows where enough fresh water seeps through to create meadows with sedges and flowering rushes such as Juncus leseurii with its bronze petals, raspberry stigmas, and creamy stamens; and finally to a mature forest of Sitka spruce. A rich understory in the spruce forest includes twinberry (Lonicera involucrata), several ferns, the calypso orchid {Calypso bulbosa), and false lily-of- the-valley (Maianthemum dilatatum), in addition to the plants found under the beach pines near Samoa. The line between the forest and the unstable dunes is con- stantly shifting, with the dunes invading the forest and then withdrawing. We'll see a lot more of the spruces up the coast. 20 Patrick's Point and Big Lagoon Continue some twenty-five miles north of Areata on State Highway 101 and follow the signs for Patrick's Point State Park. Here is a place as enchanting as Point Lobos in Monterey County, with jagged promontories and wind-swept forests. Because of heavy winds, spe- cies of the north coastal coniferous forest — a great belt of trees found in coastal Oregon, Washington, and Brit- ish Columbia—replace the redwood forest. Sitka spruce (Picea sitchensis), their western sides buffeted and trimmed by winds, grow in pure stands or are mixed with grand firs (Abiesgrandis), western hemlocks (Tsugahet- erophylla), and Douglas-fir (Pseudotsuga menziesii). In boggy spots you can find a giant, tropical-looking skunk cabbage (Lysichiton americanum) with its showy yellow flowers. Tree branches carry heavy loads of moss and epiphytic ferns such as the leather fern (Polypodium scouleri). Here also, Bishop and beach pines vie for space along the bluff edges, the bushy, light green needles of the closed-cone Bishop pine contrasting strik- ingly with the short, dark green needles of the beach pine. This is the northernmost post for the relict Bishop pine. Bishop and Monterey pines are closely related, both adapted to reproduce after fire by their whorls of permanently retained, tightly closed seed cones, whose scales peel apart when exposed to high heat. Sealed inside are thousands of seeds awaiting delivery by the natural cycle of lightning-caused fire. For this reason, stands of these pines are almost always of one age and have an uncanny uniformity as forest canopy. Gentiana sceptrum flowers growing in the bog at Big Lagoon. The jagged promontories and wind-swept forests at Patrick's Point create spectacular views. Just north of Patrick's Point is the turnoff to Big Lagoon County Park (you may walk to it along the beach), a place to see more stands of Sitka spruce, this time in combination with a sphagnum bog. Such bogs, uncommon in most of California, represent a tempo- rary successional stage like the mountain meadow, which eventually will become forest. Bordering the bog and the lagoon, spruce grows so thick that there is vir- tually no undergrowth. Under the wind-swept canopy of the spruce forest you enter another world with deep cushions of needles; old, bare branches reach out on every side, attached to the massive boles of scaly bark of fully mature trees. These spruces are immediately recognizable by their thick, spirally arranged needles with prickly tips and the clusters of long, tapered, hang- ing seed cones with fluted scales. In order to reach the bog you must first pass through the spruce forest and find an opening in a dense border of shrubs on the east side of the road. This shrubby border of salal, huckleberry, wax myrtle, and spiraea grows on higher ground and forms a nearly impenetra- ble barrier between bog and forest. In time it will be dis- placed by new generations of spruce as the margins of the pond shrink with successional changes. The bog, formed by year-round water supply from a stream fanning out before emptying into the ocean, has a rich and unusual flora worth the time to stop and see: king's gentian (Gentiana sceptrum), sneezeweed (Helenium bigelovif), tinker's penny (Hypericum anagal- loides), deer fern (Blechnum spicant), rein orchid (Habenaria dilatata), the insectivorous sundew (Drosera rotundifolia), and the rare, primitive vascular plant called running clubmoss (Lycopodium inundatum) that is unknown anywhere else in California. Redwood National Park and Prairie Creek State Park Continuing north along State Highway 101 to Orick, you pass the visitor's center for Redwood National Park, where you can take a shuttle to the Tall Trees Grove. The shuttle runs all summer on the hour and is the only easy access to the world's tallest trees, many of which grow taller than 300 feet. A more intimate way to approach the grove is to backpack up Redwood Creek from the trail head just off Bald Hills Road. The round trip is about eighteen miles. Either way you pass through fine stands of virgin redwood and a riparian forest that fol- lows Redwood Creek. There is distressing drama seen from the shuttle as you pass through the havoc wreaked upon the hills above the park by intensive logging and subsequent severe erosion. Eroded soil and rocks end up in the bed of Redwood Creek, thereby raising its water 21 level and ultimately threatening to drown the roots of redwood trees presumably already "saved" within the park boundaries. The shuttle stops just a mile above the Tall Trees Grove, and a gentle downhill hike takes you through a mixed conifer forest: Port Orford cedar {Chamaecyparis lawsoniana) with its beautifully drooping branches whose tiny scales and twiggy side branches create fans of fern-like foliage; coast hemlock with its drooping side branches, soft, light green needles, tiny seed cones, and deeply furrowed bark; and Douglas-fir with its more formal arrays of branches, lemony-scented, narrow, spirally arranged needles, and peculiar nodding seed cones with the curious three-pronged bracts extended between the scales. At the base of the slope you enter the grove itself, where a loop trail takes you through the tallest of all known trees. The trunks of these trees are arrow-straight with the first limbs beginning a hundred feet above the ground. Old crowns, where you can see them, show great age and character, often associated with the giant sequoias of the Sierra. The lush growth of a forest-edge understory attracts attention: large, spreading mountain dogwood (Cornus nuttallii) with saucers of white flower clusters; dense stands of red alder (Alnus rubra) with their feet in water; California rose-bay rhododendron (Rhododendron macrophyllum) with superb trusses of pinkish purple flowers in late May; and giant big-leaf maples (Acer macrophyllum), their canopies a tracery 22 of bright green. Ferns, shade-tolerant herbs, and mosses are everywhere, while here and there clumps of the giant Indian rhubarb (Darmera peltatum) emerge between streamside boulders. Prairie Creek State Park is to the north, with the main entrance just west of State Highway 101. This is another area of grand coastal forest. Sections along the highway have fine stands of coast redwood and some of the most lush undergrowth to be found anywhere —giant sword fern (Polystichum munitum), Trillium spp., false Solo- mon's seal (Smilacina spp.), huckelberry, redwood sorrel (Oxalis oregana), wild ginger (Asarum caudatum), inside-out flower (Vancouveria spp.), anemone (Anem- one deltoidea), and Columbia lily (Lilium columbia- num), to name a few. You can hike a beautiful loop trail from the camp- ground to the beach, passing through solemn stands of huge Sitka spruce with coast hemlock and grand fir and ending in Fern Canyon—a narrow, cool, shaded coastal canyon where sheer walls drip water all year. These walls support thousands of five-finger ferns (Adiantum peda- tum) and polypody ferns. From the trail you can see lady fern (Athryium felix-femina) and the lacy, spiny wood fern (Dryopteris dilatata) as well as the rare lily relative, twisted stalk (Streptopus amplexifolius) and maple- leafed stink currant (Ribes bracteosum). Prairie Creek is also home to a fine herd of Roosevelt elk; you can see them grazing the meadows by the campground at dawn. Left: The cones of Sitka spruce produce seeds that have long, slender wings. Bottom left: The large, pink flowers of Rhododendron macrophyllum attract many visitors to Redwood National Park and Prairie Creek State Park in late May. Below: A riparian woodland and redwood grove along the floodplain at Redwood Creek in Tall Trees Grove. Sword fern {Polystichum munitum) adds to the understory of a mature stand of redwoods. The Coastal Trail and Klamath River Just after crossing the Klamath River, turn left into the town of Requa and follow signs to the Coastal Trail. Here you can enjoy a pleasant interlude walking an easy trail across a stretch of rugged cliffs to False Klamath Cove, a distance of about three miles. You start by wind- ing through grassy fields with sweeping views of the coastline and the often fog-shrouded mouth of the Klamath. The trail quickly enters dense brush and north coastal coniferous forest with its mix of Sitka spruce, grand fir, Douglas-fir, and coast hemlock. It passes through long, tunnel-like aisles carved through hedges of prolific salmonberry (Rubus spectabilis), then follows along fern-covered embankments and shaded rills with piggyback plant (Tolmiea menziesii), western bleeding heart (Dicentra formosa), and mitrewort (Mitella ovalis). As the trail breaks out of the forest, there are broad views of rocky promontories with crashing ocean waves, hedges of cream bush (Holodiscus discolor) with its frothy cream-colored spirea-like flowers, and patches of bright rose-purple fireweed (Epilobium angustifolium). The coastal terrace prairies, probably maintained as prairies by grazing pressures, are home to several bright wildflowers, including the common daisy {Erigeron philadelphicus), yarrow (Achillea borealis), and the curi- ous, colorful firecracker flower (Dichelostemma ida- maid) with its nodding umbels of brilliant scarlet, tube- shaped flowers tipped with green recurved petals and creamy stamens. The occasional hybrid between the fire- cracker flower and ookow (Dichelostemma congestum), D. venustum, appears here with its fat,.tubular, rose- purple flowers. Finally, the walk ends at a coastal A colou> ul Ahchiiiiic rlvliimlcii in lull State Park. Krnci .11 I'laiiic Creek bog/pond called False Klamath Cove, where the yellow pond lily (Nuphar polysepalum) grows in proximity to the bog-loving Potentillapalustris with its curious red- purple flowers. Klamath Cove, which looks as if it ought to be the mouth of a river, was originally mistaken for the entrance to the Klamath River. Jedediah Smith State Park and the Smith River Just north of Crescent City, turn onto State Highway 199 and you pass through one last superb example of a redwood forest, in Jedediah Smith State Park. Other conifers sometimes grow with the redwoods, as do stands of rhododendrons and many kinds of berry bushes (thimbleberry, evergreen and deciduous huckle-