Recovery Plan

 for Coastal Plants of the 

Northern San Francisco Peninsula

Region 1

U.S. Fish and Wildlife Service

Portland, Oregon 
i

DISCLAIMER

Recovery plans delineate reasonable actions that are believed to be required to

recover and/or protect listed species. We, the U.S. Fish and Wildlife Service,

publish recovery plans, sometimes preparing them with the assistance of recovery

teams, contractors, State agencies, and others. Objectives of the plan will be

attained and any necessary funds made available subject to budgetary and other

constraints affecting the parties involved, as well as the need to address other

priorities. Recovery plans do not obligate other parties to undertake specific

actions and may not represent the views or official positions or approval of any

individuals or agencies involved in the recovery plan other than our own. They

represent our official position only after they have been signed by the Director,

Regional Director, or Manager as approved. Approved recovery plans are subject

to modification as dictated by new findings, changes in species status, and the

completion of recovery actions.

Literature Citation Should Read As Follows:

U.S. Fish and Wildlife Service. 2003. Recovery Plan for Coastal Plants of the

Northern San Francisco Peninsula. U.S. Fish and Wildlife Service, Portland,

Oregon. xvi + 304 pp.

Additional copies may be purchased from:

Fish and Wildlife Reference Service

5430 Grosvenor Lane, Suite 110 

Bethesda, Maryland 20814-2142

301-492-6403 or 1-800-582-3421; FAX: 301-564-4059

E-mail: fwrs@mail.fws.gov

http://fa.r9.fws.gov/r9fwrs

The fee for the recovery plan varies depending on the number of pages in the

plan.

An electronic version of this recovery plan will also be made available at 

http://pacific.fws.gov/ecoservices/endangered/recovery/default.htm and

http://endangered.fws.gov/recovery/index.html#plans.
ii

 ACKNOWLEDGMENTS

The draft recovery plan was prepared at the Sacramento Fish and Wildlife Office

by Peter Baye, and edited by Karen Miller and Diane Elam. The final recovery

plan was prepared by Larry Host, Kelly Hornaday, and Kirsten Tarp, and edited

by Ray Moranz, Harry McQuillen, and Don Hankins.

Marc Albert and Sharon Farrell of the Golden Gate National Recreation Area

(National Park Service) provided valuable information and insights on the past

and prospective vegetation management of the Presidio and Fort Funston, as well

as unpublished data on San Francisco lessingia and Raven’s manzanita. Joe

Cannon and Marc Albert of the Golden Gate National Recreation Area

contributed detailed site-specific information on the management and restoration

of remnant dunes supporting San Francisco lessingia. Adam Whelchel of San

Francisco State University contributed original unpublished data on the insect

visitors of San Francisco lessingia. Staci Markos of the University of California

at Berkeley contributed insights on the taxonomy of San Francisco lessingia. 

Mike Vasey of San Francisco State University made available new data and

analysis prepared with co-authors Staci Markos, Lena Hileman, and Tom Parker

(unpublished at the time earlier drafts of this recovery plan were prepared) on the

taxonomy of Raven’s manzanita. Susan Smith of San Francisco provided

information and guidance on the history and future of San Francisco lessingia’s

conservation, as well as unpublished notes on plant community composition of

lessingia sites. We also acknowledge the valuable recent scientific contributions

of Yolanda Mollette and Crima Pogge, for their studies of San Francisco dune

seed banks and lessingia population ecology, respectively. Sylvia Martin of the

U.S. Fish and Wildlife Service graciously assisted preparation of photographic

reproductions.

We are grateful to the peer reviewers who provided us with their comments and

suggestions. Their contribution was essential to making the plan scientifically

sound.

We take responsibility for any errors that remain, as we may not have grasped all

the nuances of the information provided to us, despite a sincere desire to blend in

all of the improvements communicated through the many comments received.
iii

EXECUTIVE SUMMARY

Current Species Status: This recovery plan features two plant species, both of

which are federally listed as endangered and are endemic to the northern San

Francisco peninsula in California. San Francisco lessingia (Lessingia

germanorum), an annual herb in the aster family, exists at six sites in the Presidio

of San Francisco and one site in Daly City. It is threatened by invasion of

nonnative vegetation, habitat loss, and potential land use conflicts in limited

restorable urban habitat. Raven’s manzanita (Arctostaphylos hookeri ssp. ravenii)

is a rare evergreen creeping shrub in the heath family. Only one genetic

individual exists in the wild, but some clones of the original and some of its

seedlings have been artificially propagated. The original plant is located in

remnant coastal scrub and grassland in the Presidio, along with its transplanted

clones. It is threatened by failure to reproduce naturally, habitat loss, potential

land use conflicts in limited restorable urban habitat, and disease.

Geographic and Ecological Scope of the Recovery Plan: The ecosystems on

which the endangered species of this recovery plan depend are limited to narrow

geographic areas of the northern San Francisco peninsula. Their habitats are

further restricted to specific substrate types: old coastal sand deposits (San

Francisco lessingia) and outcrops of serpentine or similar rocks (Raven’s

manzanita). Both taxa grow only in sparse, relatively open native dune scrub,

coastal scrub, and grassland vegetation.

Taxonomic Scope of the Recovery Plan: This recovery plan serves as a

revision for the Raven’s Manzanita Recovery Plan (U.S. Fish and Wildlife

Service 1984), addressing Raven’s manzanita jointly with San Francisco lessingia

to facilitate a more holistic ecosystem-based approach for these species. This

recovery plan considers actions for the two featured endangered species along

with conservation needs of a number of plants and animals that are associated

with them. Several other associated federally-listed taxa that are already covered

by other recovery plans are discussed herein: bay checkerspot butterfly

(Euphydryas editha bayensis), Marin dwarf-flax (Hesperolinon congestum), and

Presidio clarkia (Clarkia franciscana), covered by the Recovery Plan for

Serpentine Soil Species of the San Francisco Bay Area (U.S. Fish and Wildlife
iv

Service 1998a); and Myrtle’s silverspot butterfly (Speyeria zerene myrtleae) and

beach layia (Layia carnosa), covered by the Recovery Plan for Seven Coastal

Plants and the Myrtle’s Silverspot Butterfly (U.S. Fish and Wildlife Service

1998b). In addition, the recovery plan considers 16 plant species of concern

associated with San Francisco lessingia and Raven’s manzanita. These 16 plant

species are known to be at risk of either range collapse (local or regional

extinction) or range-wide extinction. Seventeen other plant species of local or

regional conservation significance are also considered. These plants share

habitats with the listed species and have suffered substantial declines on the

northern San Francisco Peninsula and adjacent coast.

Recovery Priority: San Francisco lessingia: 2C (full species; high threats;

high recovery potential; conflict with development

projects)

Raven’s manzanita: 12 (subspecies; moderate threats; low

recovery potential)

Recovery priority numbers are based on criteria published by Federal Register

Notice (U.S. Fish and Wildlife Service 1983a, 1983b).

Recovery Objectives: The objectives of this recovery plan are to conserve and

restore sufficient habitat and populations of San Francisco lessingia and Raven’s

manzanita to warrant their reclassification from endangered to threatened, and

ultimately to delist San Francisco lessingia. Recovery of Raven’s manzanita

sufficient to warrant delisting is not projected for the foreseeable future.

Actions Needed: This recovery plan does not focus on actions that only benefit

managed populations of the two listed taxa featured in this plan. Instead, it

stresses re-establishing dynamic, persistent populations of the listed taxa within

plant communities that have been restored so they are as “self-sustaining” as

possible within urban wildland reserves. In self-sustaining communities there is

the insurance of replacement in perpetuity of individuals in the population

through natural reproduction. Recovery actions for San Francisco lessingia focus

on the restoration and management of large, dynamic mosaics of coastal dune
v

areas supporting shifting populations within the species’ narrow historic range. 

Recovery of Raven’s manzanita will include, but will not be limited to, the

original recovery strategy presented in the Raven’s Manzanita Recovery Plan

(U.S. Fish and Wildlife Service 1984). This strategy emphasized the stabilization

of the single remaining genetic individual, which is necessary, but without further

measures would result in a dead end to the species’ evolution. This recovery plan

seeks to re-establish multiple sexually reproducing populations of Raven’s

manzanita in association with historically associated species of local serpentine

outcrops. 

We recognize that the implementation of recovery plans is always affected by

circumstances and new information. Furthermore, this recovery plan focuses on

biological analysis; review of other aspects of implementing the recovery actions

in this plan will be done as required by such laws as the National Environmental

Policy Act and the National Historic Preservation Act. If requested funding is not

approved, and other partnerships cannot be formed, the implementation schedule

will have to be adjusted to the means available.

Specific actions needed include:

(1) Protect and restore a series of ecological urban wildland reserves.

(2) Promote population increases of target species within urban wildland reserves

and reintroduce target species to restored habitat.

(3) Long-term removal (local eradication) or suppression of invasive, nonnative

vegetation within and around all reserves and subsequent reestablishment of

native communities compatible with endangered species within the ecological

reserves.

Recovery actions will occur in three identified recovery units for San Francisco

lessingia including: (1) the Presidio of San Francisco (National Park Service and

Presidio Trust lands); (2) Fort Funston (National Park Service lands) and Hillside

Park (City of Daly City); and (3) the satellite reserves, which are smaller urban

park dune remnants (City of San Francisco and National Park Service lands). 
vi

Actions for Raven’s manzanita will occur in serpentine bedrock and soil outcrops

of the Presidio, mostly along bluffs of the north shore, and on hilltop bedrock

outcrops at selected locations within San Francisco (city and some Federal lands).

Recovery Criteria:

San Francisco lessingia will be considered for downlisting to threatened status

when interim recovery criteria are met and the Lobos Dunes unit has expanded to

Battery Caulfield Road and upper Baker Beach. The species will be considered

for delisting when long-term recovery criteria are met.

Interim Recovery Criteria

(1) Long-term expansion of existing populations and reduction of

nonnative vegetation occurs in dune reserves in the Presidio Recovery

Unit (Lobos Creek, Battery Caulfield, Wherry, Rob Hill, and Public

Health Services Hospital sites). The populations in these reserves are

expected to fluctuate but should not decline below 50,000, 1,000, 5,000,

5,000, and 5,000, respectively. Cover of nonnative vegetation in these

reserves should be less than 5 percent, 20 percent, 5 percent, 20 percent,

and 20 percent, respectively. Research and monitoring are expected to

determine the most efficient methods for control of nonnative vegetation

and may result in modification of these recovery criteria.

(2) The population of the Daly City reserve shows no net long-term

decrease. Populations are expected to fluctuate but should not decline

below 50,000 plants. Cover of nonnative vegetation should show no

progressive increase over more than 2 years.

(3) Recreation management needs for maintenance of a dune scrub

community in the Fort Funston Reserve are established through sitespecific studies, and a reintroduced population has persisted over a full

precipitation cycle. The population is expected to fluctuate but should

achieve a minimum self-sustaining population size of approximately

500,000 plants within the first ten years after founders are transplanted.
vii

(4) At least 1,000 seeds representing both the existing Presidio and Daly

City populations are stored and maintained in qualified botanical gardens

as insurance against extinction in the wild. 

Long-term Recovery Criteria

All reserves must be protected in perpetuity with appropriate vegetation

management.

(1) Expanded, restored reserves with natural vegetation and dune

dynamics are established in the Presidio Recovery Unit. The area

including Baker Beach dunes, Lobos Dunes and nearby conifer groves,

Wherry Dunes and Housing sites, and the Battery Caulfield Road site

must be restored to a contiguous dune field (approximately 44 hectares

[110 acres]) with unobstructed wind fetch to the Golden Gate, locally

steep dune slopes, and a natural successional mosaic of active and

stabilizing dune blowouts (population at least 500,000 plants; nonnative

vegetative cover must not exceed 5 percent during the first 10 years of

restoration and must decline over the first 15 years). Dune habitat at the

Rob Hill reserve area must increase to 2 hectares (5 acres) and the

southwest slope of Rob Hill must be restored to dune scrub (population at

least 100,000 plants; nonnative vegetative cover must not exceed 5

percent). At least 3 hectares (7 acres) of the Public Health Services

Hospital dune area above the slope must be restored to native dune

vegetation (population at least 50,000 plants; nonnative vegetative cover

must not exceed 5 percent). Research and monitoring are expected to

determine the most efficient methods for control of nonnative vegetation

and may result in modification of these recovery criteria.

(2) At least 1.2 hectares (3 acres) of Daly City Reserve in the Southern

Recovery Unit are cleared of nonnative vegetation and intensively

managed (population at least 50,000 plants; no increases in nonnative

vegetation).

(3) Dune restoration and vegetation management should be done on 30

hectares (75 acres) at Fort Funston Reserve in the Southern Recovery Unit
viii

sufficient to establish ecosystem function and support a self-sustaining

population of San Francisco lessingia. Site-specific studies of recreational

use management compatible with maintenance of a dune scrub community

must be completed to further define the restoration and management

strategies. Buffers of landscape vegetation around all historic structures

and buffer management needs will also need to be identified. A

population should be reintroduced from the Daly City seed source

(population at least 10,000 plants after 5 years; must reach 500,000 plants

after 10 years; new colonies must spontaneously establish within 10

years). Nonnative woody vegetative cover must be below 1 percent;

iceplant and European beachgrass cover must decline and be below 10

percent after 10 years. Research and monitoring are expected to

determine the most efficient methods for control of nonnative vegetation

and may result in modification of these recovery criteria.

(4) Populations must be introduced in the Satellite Recovery Unit (should

reach 100,000 plants within 10 years, with minimum size of 5,000 plants).

This criterion is preliminary subject to additional information. The

locations of dune areas suitable for management within this Recovery Unit

are scattered around the western half of San Francisco County. The

specific size, number, and configuration of reserves needs to be

determined on the basis of more detailed site-specific information.

Raven’s manzanita will be considered for downlisting to threatened when interim

recovery criteria are met, five spontaneously reproducing, ecologically,

morphologically, and genetically variable populations are established in reserves

in San Francisco outside the Presidio, two sexually reproduced generations are

established within the Presidio, and net population size (net number of clones)

and net individual mean (average) clone size increase at all sites over 30 years.

Interim Recovery Criteria

(1) The original site of the remnant clone and the sites of its daughter

clones are maintained and protected in perpetuity. All of the clones in the

wild must increase in size over a 10-year period, and the number of clones

must increase in the same period.
ix

(2) Cultivated populations of Raven’s manzanita are perpetually

maintained at two or more botanical gardens. Populations must include 50

daughter clones of the original Presidio plant, with an additional goal of

50 seedling-grown plants with at least two clonal replicates each. 

(3) Five or more spontaneously reproducing new populations (comprising

at least five daughter clones each) are established in reserves on bedrock

outcrops in San Francisco outside the Presidio, preferably in proximity to

historic localities. At least three reserves must be on serpentine substrates.

New colonies must show net growth 5 years after transplanting with

intensive maintenance and 5 years after cessation of maintenance.

(4) The taxonomic relationships and reproductive biology of Raven’s

manzanita are studied.

Estimated Costs of Recovery: Approximately $23,432,500 (plus costs to be

determined). Costs to be determined include: the costs of public outreach, the

costs of implementing vegetation management plans, habitat acquisition for

Raven’s manzanita, and site preparation for Raven’s manzanita. 

Date of Recovery:

Raven’s manzanita: If recovery criteria are met, downlisting to threatened could

occur by 2030.

San Francisco lessingia: If recovery criteria are met, downlisting to threatened

could occur by 2020, and delisting could occur by 2030. 
x

TABLE OF CONTENTS

I. INTRODUCTION ...............................................1

A. Scope and Purpose of the Recovery Plan . . . . . . . . . . . . . . . . . . . . . . 1

B. Coastal Dunes and Bedrock Outcrops of the Northern San Francisco

Peninsula: Physical and Ecological Background .............4

1. The San Francisco Dune System . . . . . . . . . . . . . . . . . . . . . . . 4

a. Geomorphology and Evolution of the San Francisco

Dune System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

b. Native Vegetation and Flora of the San Francisco Dune

System. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

c. Dynamics of Dune Vegetation . . . . . . . . . . . . . . . . . . 14

2. San Franciscan Bedrock Outcrops and their Vegetation . . . . 15

a. San Francisco Bedrock Outcrops . . . . . . . . . . . . . . . . 15

b. Vegetation of San Francisco Bedrock Outcrops . . . . . 19

II. BIOLOGY OF FEDERALLY LISTED SPECIES . . . . . . . . . . . . . . . . . . . . 21

A. San Francisco Lessingia (Lessingia germanorum) . . . . . . . . . . . . . . 21

1. Description and Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

2. Historic and Current Distribution and Abundance . . . . . . . . . 24

a. Modern Presidio Populations . . . . . . . . . . . . . . . . . . . 24

b. Southwestern San Francisco and Vicinity Populations

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

c. Variation in Abundance of San Francisco Lessingia

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

3. Ecology and Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

a. Environmental Conditions . . . . . . . . . . . . . . . . . . . . . 37

b. Plant Associations . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

c. Life History and Reproduction . . . . . . . . . . . . . . . . . . 39

4. Causes of Decline and Threats to Survival . . . . . . . . . . . . . . . 42

5. Conservation Efforts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

6. Species Recovery Strategy for San Francisco Lessingia . . . . 51

B. Raven’s Manzanita (Arctostaphylos hookeri ssp. ravenii) . . . . . . . . 55

1. Description and Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
xi

2. Current and Historic Distribution . . . . . . . . . . . . . . . . . . . . . . 61

3. Ecology and Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . 63

4. Causes of Decline and Threats to Survival . . . . . . . . . . . . . . . 68

5. Conservation Efforts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

6. Species Recovery Strategy for Raven’s Manzanita . . . . . . . . 75

III. OTHER FEDERALLY LISTED SPECIES AND SPECIES OF CONCERN

OR REGIONAL CONSERVATION SIGNIFICANCE . . . . . . . . . . . . . 81

A. Ecologically Associated Federal Listed Species . . . . . . . . . . . . . . . . 82

1. Presidio Clarkia, Clarkia franciscana . . . . . . . . . . . . . . . . . . . 82

2. Marin Dwarf-flax, Hesperolinon congestum . . . . . . . . . . . . . . 89

3. Beach Layia, Layia carnosa . . . . . . . . . . . . . . . . . . . . . . . . . . 89

4. Myrtle’s Silverspot Butterfly, Speyeria zerene myrtleae . . . . 90

5. Bay Checkerspot Butterfly, Euphydryas editha bayensis . . . . 91

B. Species of Concern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91

1. Franciscan Manzanita, Arctostaphylos franciscana . . . . . . . . 95

2. Franciscan Thistle, Cirsium andrewsii . . . . . . . . . . . . . . . . . . 96

3. San Francisco Spineflower, Chorizanthe cuspidata . . . . . . . . 98

4. Round-headed Chinese Houses, Collinsia corymbosa . . . . . . 98

5. San Francisco Wallflower, Erysimum franciscanum . . . . . . . 99

6. San Francisco Gumplant, Grindelia hirsutula var. maritima 100

7. Diablo Helianthella, Helianthella castanea . . . . . . . . . . . . . 100

8. Evax, Hesperevax sparsiflora . . . . . . . . . . . . . . . . . . . . . . . . 101

9. Kellogg’s Horkelia, Horkelia cuneata ssp. sericea

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

10. Large-flowered Linanthus, Linanthus grandiflorus . . . . . . 102

11. Curly-leaved Monardella, Monardella undulata . . . . . . . . 102

12. Greene’s Popcornflower, Plagiobothrys reticulatus var.

rossianorum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

13. San Francisco Campion, Silene verecunda ssp. verecunda 104

14. Seashore or Coast Starwort, Stellaria littoralis . . . . . . . . . 104

15. Dune Tansy, Tanacetum camphoratum . . . . . . . . . . . . . . . 105

16. San Francisco Owl’s-clover, Triphysaria floribunda . . . . . 106

C. Species of Local and Regional Conservation Significance . . . . . . . 106
xii

1. Pink Sand-verbena (introgressant) Abronia umbellata ssp.

umbellata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107

2. Coast Rock-cress, Arabis blepharophylla . . . . . . . . . . . . . . 109

3. Nuttall’s Milk-vetch, Astragalus nuttallii var. virgatus . . . . 109

4. California Saltbush, Atriplex californica . . . . . . . . . . . . . . . 110

5. Indian Paintbrush, Castilleja affinis ssp. affinis

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111

6. Banded (Broadleaf) Purple Owl’s-clover, Castilleja exserta

ssp. latifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

7. California Goosefoot, Chenopodium californicum . . . . . . . . 114

8. Davy’s Clarkia, Clarkia davyi . . . . . . . . . . . . . . . . . . . . . . . 115

9. California Croton, Croton californicus . . . . . . . . . . . . . . . . . 115

10. Franciscan Leafy-daisy, Erigeron foliosus var. franciscensis

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115

11. Yarrow-leaf Gilia, Gilia millefoliata . . . . . . . . . . . . . . . . . 116

12. San Francisco (Chamisso’s) Dune Gilia, Gilia capitata ssp.

chamissonis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

13. Wedge-leaf Horkelia, Horkelia cuneata ssp. cuneata . . . . 117

14. Common Linanthus, Linanthus parviflorus . . . . . . . . . . . . 117

15. Skunkweed, Navarretia squarrosa . . . . . . . . . . . . . . . . . . . 118

16. California Broomrape, Orobanche californica ssp. californica

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118

17. Coast Rein-orchid or Coast Piperia, Piperia elegans . . . . . 118

IV. RECOVERY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

A. Objectives and Criteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

1. Recovery Criteria for San Francisco Lessingia . . . . . . . . . . 121

a. Interim Recovery Criteria . . . . . . . . . . . . . . . . . . . . . 127

i. Presidio Recovery Unit . . . . . . . . . . . . . . . . . 127

ii. Daly City Reserve. . . . . . . . . . . . . . . . . . . . . 132

iii. Fort Funston Reserve. . . . . . . . . . . . . . . . . . 132

iv. Offsite. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133

b. Long-term Recovery Criteria . . . . . . . . . . . . . . . . . . 133

i. Presidio Recovery Unit . . . . . . . . . . . . . . . . 134

ii. Southern Recovery Unit . . . . . . . . . . . . . . . . 140
xiii

iii. Satellite Recovery Unit . . . . . . . . . . . . . . . . 144

2. Recovery Criteria for Raven’s Manzanita . . . . . . . . . . . . . . 146

a. Interim Recovery Criteria . . . . . . . . . . . . . . . . . . . . . 147

i. Habitat and Population Stabilization in the

Presidio . . . . . . . . . . . . . . . . . . . . . . . . . 147

ii. Propagation of Seedling and Clonal Stock . . 147

iii. Establishment of New Daughter Clones on

Presidio Serpentine Bluff Sites . . . . . . . 148

iv. Investigation of Taxonomic Relationships and

Reproduction . . . . . . . . . . . . . . . . . . . . . 148

b. Long-term Recovery Criteria . . . . . . . . . . . . . . . . . . 149

i. Reproduction and Growth in the Presidio Bluff

Population . . . . . . . . . . . . . . . . . . . . . . . 149

ii. Establishment and Protection of New Interior

Populations . . . . . . . . . . . . . . . . . . . . . . . 149

iii. Permanent Reserve Cultivated Populations in

Botanical Gardens . . . . . . . . . . . . . . . . . 150

B. Comprehensive Strategy of Recovery Actions . . . . . . . . . . . . . . . . 150

1. Recovery Strategy for San Francisco Lessingia . . . . . . . . . . 150

a. Presidio Recovery Unit . . . . . . . . . . . . . . . . . . . . . . . 151

i. General Requirements of the Presidio Recovery

Unit . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

ii. Site-specific Requirements of the Presidio

Recovery Unit . . . . . . . . . . . . . . . . . . . . 154

b. Southern Recovery Unit . . . . . . . . . . . . . . . . . . . . . . 162

i. Daly City Reserve . . . . . . . . . . . . . . . . . . . . . 162

ii. Fort Funston Reserve . . . . . . . . . . . . . . . . . . 164

c. Satellite Recovery Unit . . . . . . . . . . . . . . . . . . . . . . . 167

i. Sutro Heights Sub-Unit . . . . . . . . . . . . . . . . . 167

ii. Sunset Heights Sub-Unit . . . . . . . . . . . . . . . 168

2. Recovery Strategy for Raven’s Manzanita . . . . . . . . . . . . . . 170

a. Development of a New “Pure” Inbred Founder

Population of Raven’s manzanita . . . . . . . . . . . 172

b. Contingency Measures for Producing New Individuals:

Introgressive Breeding of Raven’s Manzanita . . 173
xiv

c. Site Selection and Acquisition for New Populations in

San Francisco . . . . . . . . . . . . . . . . . . . . . . . . . . . 176

d. Control of Cultivated Manzanita Pollen Flow and

Spontaneous “Nonnative” Hybrids . . . . . . . . . . 177

e. Identification and Permanent Protection of Existing and

Potential Serpentine Habitat in the Presidio . . . 178

f. Public Outreach and Education . . . . . . . . . . . . . . . . . 178

g. Additional Research . . . . . . . . . . . . . . . . . . . . . . . . . 179

3. Conservation Recommendations . . . . . . . . . . . . . . . . . . . . . 179

V. STEPDOWN NARRATIVE OF RECOVERY ACTIONS . . . . . . . . . . . . . 181

VI. IMPLEMENTATION SCHEDULE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205

VII. REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218

A. Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218

B. Personal Communications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236

C. Unpublished References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 238

D. In Litt. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239

VIII. APPENDICES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I-1

APPENDIX I. Native Vascular Plant Species of Coastal Dunes and Dune

Slacks of the Northern San Francisco Peninsula . . . . . . . . . . . . I-1

APPENDIX II. Native Vascular Plant Species of Serpentine Outcrops 

of the Northern San Francisco Peninsula . . . . . . . . . . . . . . . . . II-1

APPENDIX III. Invasive Nonnative Plants That May Affect Recovery of 

San Francisco Lessingia and Raven’s Manzanita . . . . . . . . . . III-1

APPENDIX IV. Glossary of Technical Terms . . . . . . . . . . . . . . . . . . IV-1

APPENDIX V. Summary of Threats and Recommended Recovery

Actions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . V-1

APPENDIX VI. Agency and Public Comments on the Draft Recovery

Plan for the Coastal Plants of the Northern San Francisco

Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VI-1
xv

LIST OF TABLES

Table 1. Historic locality and habitat data from herbarium specimens of San

Francisco lessingia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

LIST OF FIGURES

Figure 1. Approximate distribution of San Francisco dune sheet, and

approximate modern and extirpated localities of San Francisco

lessingia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Figure 2. Localities of serpentine and greenstone bedrock outcrops in San

Francisco.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Figure 3. Geological control of Raven’s manzanita habitat . . . . . . . . . . . . . . 18

Figure 4. San Francisco lessingia (Lessingia germanorum) . . . . . . . . . . . . . . 22

Figure 5. Approximate contemporary locations of San Francisco lessingia in

the Presidio, San Francisco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Figure 6. Approximate contemporary location of San Francisco lessingia at

Hillside Park, Daly City . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Figure 7. Distribution of recovery units for San Francisco lessingia on the San

Francisco Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

Figure 8. Raven’s manzanita (Arctostaphylos hookeri ssp. ravenii) . . . . . . . . 56

Figure 9. Selected plant species from the northern San Francisco Peninsula. 83

Figure 10. Presidio Recovery Unit for San Francisco lessingia . . . . . . . . . . . 129
xvi

Figure 11. Presidio Recovery Unit. Area detail: proposed reintroduction,

restoration, and protection areas within the Presidio . . . . . . . . . . . 130

Figure 12. Southern Recovery Unit for San Francisco lessingia. Proposed

reintroduction, restoration, and protection areas within Fort Funston

dunes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
1

I. INTRODUCTION

A. Scope and Purpose of the Recovery Plan

The two federally listed plant species addressed in this recovery plan are endemic

to the San Francisco Peninsula: San Francisco lessingia (Lessingia germanorum

Cham.), an annual herb of older sand dunes and similar sparsely vegetated coastal

sand deposits (Lane 1993); and Raven’s manzanita (recently treated

taxonomically as Arctostaphylos hookeri G. Don ssp. ravenii P. Wells), a

prostrate evergreen shrub found naturally only on certain types of Franciscan

formation bedrock outcrops, particularly serpentine (U.S. Fish and Wildlife

Service 1984). This recovery plan merges an update of the Raven’s Manzanita

Recovery Plan (U.S. Fish and Wildlife Service 1984) with a new plan for the

recovery of San Francisco lessingia. It addresses actions that would benefit other

federally listed species that are ecologically associated with these principal

species within this geographic area, but are covered comprehensively in other

recovery plans. Other federally listed plant species, covered principally in other

recovery plans, but sharing potential habitat and geographic range, are two annual

herbs: a local serpentine endemic, Presidio clarkia (Clarkia franciscana), and the

rare but wide-ranging beach layia (Layia carnosa), restricted to coastal dunes

(U.S. Fish and Wildlife Service 1998a, 1998b).

The current geographic distributions of San Francisco lessingia and Raven’s

manzanita have been markedly reduced by habitat loss in San Francisco. 

Populations occur primarily within small, highly altered local remnants of dune

and bedrock outcrop vegetation on former military lands of the Presidio. 

Herbarium records indicate that San Francisco lessingia was a local element of

some phases of coastal dune scrub communities on the extensive sand dune sheet

of San Francisco, and are consistent with records of many historically associated

dune plants (Appendix I). It is associated with areas of sparse, low vegetative

cover in older dunes. Raven’s manzanita occurred in local abundance with a

closely related subspecies from which it had not been distinguished until recent

decades (Franciscan manzanita; Arctostaphylos franciscana, synonym =

Arctostaphylos hookeri ssp. franciscana). Both manzanitas occurred on scattered

exposures of bedrock outcrops composed of serpentine and greenstone (mafic and
2

ultramafic igneous rocks, derived from altered minerals of deep magmas rich in

heavy metals) (U.S. Fish and Wildlife Service 1984). Both manzanitas formerly

occurred together at several sites. A single individual of Raven’s manzanita

exists in the wild today, but Franciscan manzanita now exists only in cultivation.

Most of the dune ecosystem and local bedrock outcrop plant communities of San

Francisco County have been eliminated. Recovery of these listed species within

protected and restored portions of their respective ecosystems is possible,

however, and is the ultimate goal of this recovery plan. The geographic and

ecological focus of this recovery plan, therefore, is on semi-natural vegetation of

old dunes and bedrock outcrops on the few remaining undeveloped portions of the

San Francisco Peninsula. The purpose of recovery is to reinforce or reestablish

viable populations of San Francisco lessingia and Raven’s manzanita, and their

associated species of concern, in managed urban reserves of semi-natural

vegetation, incorporating natural ecological dynamics, patterns, and processes to

the greatest extent practical. These areas are found in the southwestern part of the

Presidio, lands of the Golden Gate National Recreation Area, portions of urban

parks, and on undeveloped private and municipal lands. Because of the

essentially urban land use context, recovery objectives necessarily depend on

some active vegetation management and integration with public recreational uses

of urban parklands.

Two other federally listed plant species historically occurred on the northern San

Francisco Peninsula in wetland habitats, and are treated in other recovery plans: 

California sea-blite (Suaeda californica S. Watson) (U.S. Fish and Wildlife

Service 1994a) and marsh sandwort (Arenaria paludicola Robinson) (U.S. Fish

and Wildlife Service 1993). 

California sea-blite is a gray-green shrub in the goosefoot family

(Chenopodiaceae) that is now restricted to sandy high salt marsh edges of Morro

Bay. San Francisco Bay populations of this species reported from the early 20th

century (Jepson 1911, Howell et al. 1958) are now extinct. California sea-blite is

planned for reintroduction at the Crissy Field barrier beach and salt marsh

restoration project in the Presidio. The reintroduction of this species to San

Francisco Bay salt marshes, and its recovery, are treated in the Draft Recovery
3

Plan for Tidal Marsh Ecosystems of Central and Northern California (U.S. Fish

and Wildlife Service, in prep.).

Marsh sandwort is a low-growing clonal perennial herb in the pink family

(Caryophyllaceae). This species is restricted to coastal sedge marshes, lagoon

edges, and swampy thickets at widely disjunct localities, including Black Canyon

Lake (San Luis Obispo County, the only currently verified population), Presidio

marsh (now Crissy Field, San Francisco; Brandegee 1892, Howell et al. 1958),

Santa Cruz and San Bernardino Counties, and the Tacoma area of Washington

State. It also has been reported from Mexico (U.S. Fish and Wildlife Service

1998c). It was among the first plants in San Francisco to become locally extinct

early in the city’s history (Behr 1892). Neither marsh sandwort nor California

sea-blite overlap ecologically with the dry, terrestrial, sparsely vegetated habitats

of the species covered in this plan. Marsh sandwort recovery is treated

comprehensively in the Draft Recovery Plan for Marsh Sandwort and Gambel’s

Watercress (U.S. Fish and Wildlife Service 1998c).

In addition, the coastal scrub and grassland of the San Francisco Peninsula has

historically supported habitat for the San Francisco endemic, Xerces blue

butterfly (Glaucopsyche xerces), the first butterfly known to have become extinct

in the history of the United States (Powell and Hogue 1979). The area also

supported habitat for the endangered Myrtle silverspot butterfly (Speyeria zerene

myrtleae), a species formerly associated with coastal scrub and grassland from

San Mateo to Marin Counties and currently extirpated south of the Golden Gate. 

Its recovery is addressed in the Recovery Plan for Seven Coastal Plants and

Myrtle Silverspot Butterfly (U.S. Fish and Wildlife Service 1998b). The bay

checkerspot butterfly (Euphydryas editha bayensis) is a federally threatened

subspecies associated with serpentine habitats in the Bay area; its recovery is

treated in the Recovery Plan for Serpentine Soil Species of the San Francisco Bay

Area (U.S. Fish and Wildlife Service 1998a).
4

B. Coastal Dunes and Bedrock Outcrops of the Northern San Francisco

Peninsula: Physical and Ecological Background

1. The San Francisco Dune System

a. Geomorphology and Evolution of the San Francisco Dune System. Prior to

urbanization, the San Francisco dune system was among the largest in California

(Cooper 1967). The dune sheet (a continuous mantle of wind-blown sand

covering underlying substrate) extended from Land’s End to south of Lake

Merced (8 kilometers [5 miles]), and reached across the entire Peninsula (nearly

12 kilometers [7.5 miles] wide near the north end of the Peninsula) (Schlocker

1974) (Figure 1). The dune system of San Francisco is relatively isolated: the

nearest coastal dune sheets of comparable size are at Point Reyes and Monterey

Bay. The modern isolation of the San Francisco dune system is dependent on sea

level. During periods of low sea level in the Pleistocene and early Holocene

epochs, a broad coastal plain and delta were exposed far offshore from modern

shorelines (Schlocker 1974, Atwater et al. 1979, Howard 1979). Coastal plain

shorelines typically develop barrier beach coasts that support dune systems

(Leatherman 1979, Davies 1980), so antecedents of modern central coast dune

systems may have been widespread or continuously distributed along the central

California coast during the many periods of glacial low sea levels during the

Pleistocene epoch.

The western portions of the San Francisco dune sheet were described in the early

19th century as large expanses of relatively unvegetated, desert-like sand with

wavy topography (Cooper 1967), suggesting extensive transverse dunes (dune

ridges like giant ripples, elongated perpendicular to dominant winds) similar to

those found on the dune sheet of Coos Bay in Oregon. Photographs of the “outer

lands” of San Francisco from the 1860's reveal large expanses of mobile sand

with discrete vegetated dune hummocks (G. Gaar unpublished local historic

photography 1999). Mobile, sparsely vegetated or bare transverse dunes are

evident in large undeveloped tracts of the Sunset district (south of Golden Gate

Park) in 1930's aerial photographs (U.S. Army Corps of Engineers, San Francisco

District, file information), and were observed by Schlocker in 1937 
5

Figure 1. Approximate distribution of San Francisco dune sheet with

unconsolidated surface deposits of wind-blown sand (Schlocker 1974,

Cooper 1967, and U.S. Coast survey T-sheets from the 1850's). The

relationship between the landward extent of the dune sheet and the bay

shoreline has been obscured by bay fill. Approximate modern localities of

San Francisco lessingia are shown in circled stars (k). The extirpated

historic collection localities are shown in solid black circles (). Lake

Merced and Ocean View locations are generalized and not site-specific. 
6

(Schlocker 1974). The nearest dune system persisting today with comparable but

smaller scale transverse dunes is at the outer Dillon Beach dune complex at the

mouth of Tomales Bay, Marin County. San Francisco and Dillon Beach dunes

both developed ponds and wetlands in depressions between dunes (Cooper 1967),

probably fed from dune-dammed streams, springs, and seasonally high

groundwater tables, as occurs at Dillon Beach dunes today (P. Baye unpublished

data 1997 to 1998).

More eastern and interior San Francisco dunes were described as covered with

scrubby trees (mostly coast live oak [Quercus agrifolia]) and shrubs (Howell et

al. 1958, Clary 1980), and were therefore probably much older than dunes of

“western lands” (Cooper 1967, Schlocker 1974). Local dune sheets, independent

of the main sheet that extended eastward from Ocean Beach, also were found in

association with beaches along the north shore of the Peninsula, such as Baker

Beach, the Presidio sand spit and Strawberry Island (now Crissy Field), and Black

Point (former shoreline southeast of Crissy Field) (Cooper 1967). Apparently

ancient small remnants of dune soils also occur within portions of serpentine

landslides of the Presidio bluffs (northeast of Baker Beach), Sutro Heights, and

Land’s End (P. Baye, pers. observ.), indicating former presence of wide beaches

(sand sources) and climbing dunes along what are now steep, narrow, erosional

sand and boulder beaches.

The San Francisco dunes were oriented with northwest winds, probably due to

local topographic deflection of offshore winds that otherwise tend to be more

southwesterly (Cooper 1967). Deflection of winds by local topography caused

local major anomalies in dune trends such as the north-south trending dune

tongue west of Laguna Honda, in the lee of Sunset Heights (Figure 1). The

eastern portions of the dune sheet, as well as areas sheltered by hills of Franciscan

bedrock, included longitudinal dunes and irregular partially-eroded deposits with

vegetative cover of scrub and low oaks (Clary 1980). Climbing dunes ascended

Franciscan bedrock hills (Buena Vista, Mt. Sutro, Sutro Heights, Sunset Heights,

Lone Mountain) to heights over 180 meters (600 feet) in elevation (Schlocker

1974), often thinning to sand veneers over underlying soil and bedrock. West of

Lake Merced, the dune sheet is perched on eroding marine bluffs (and buried

soils) composed mostly of weakly consolidated, uplifted sand deposits of the
7

Merced formation (Pleistocene sandy “fossil” marine and dune deposits that rise

abruptly at Fort Funston; Schocker 1974). Evidence based on sand mineralogy

and transport alongshore suggests that erosion of Merced deposits has been a

principal long-term source of sand for the San Francisco dune system (Cooper

1967, Schlocker 1974). The dune sheet also overrides the similar poorly

consolidated sand deposits of the Colma formation east of Lake Merced (Bonilla

1965).

The modern San Francisco dune system is probably derived from a composite of

earlier and later generations of dunes deposited during the Pleistocene (glacial)

epoch and recent post-glacial (Holocene) times (Schlocker 1974). Cooper (1967)

identified two discrete advances or episodes of mobility in California dune

systems, based on comprehensive comparative analysis of Pacific coast dune

systems. The earlier advance, “Episode I,” which may have corresponded with

low glacial/early post-glacial (late Pleistocene/early Holocene) sea level and wide

sandy coastal plains, corresponds with remnant landward dunes with welldeveloped chaparral, mature dune scrub and soil, grassland, or oak woodland. 

The later “Episode II” dune advance is associated with (or grades into) recent

depositional environments. The outer mobile transverse dunes of the western

parts of San Francisco were apparently “Episode II,” or later Holocene deposits. 

Urbanization has eliminated evidence of probable “Episode I” dune boundaries in

San Francisco, but these boundaries are identifiable in dune systems north and

south of San Francisco (Bodega Head, Dillon Beach, Monterey Bay; Cooper

1967). Traces of highly weathered and organic-stained old dune soils in

landslides of the Presidio bluffs, and remnant coast live oak stands on old dune

soils in Golden Gate Park may represent remnants of “Episode I” dunes. Soil

properties of older dunes affect the diversity of the plant communities they

support (Ranwell 1972). The former dunes at the east end of the City supported

oak woodland or mature dune scrub with well-developed soil profiles and

weathered organically-stained sand. This vegetation-soil unit possibly represents

“Episode I” dunes of early Holocene or late Pleistocene age. 

The dunes perched above the bluffs of the Merced formation may be derived from

mixed sources of sand by wind erosion of: (1) consolidated, weathered yellow-
8

brown Merced formation sands exposed in the marine scarp; (2) loose,

unconsolidated sand of slope failures in Merced deposits; and (3) unweathered

gray-white sand transported directly from the beach. The development of new

dunes from reworked ancient sand from bluffs, sometimes mixed with recent

sand, was described for the former Laguna Salada dunes located to the south of

San Francisco (Cooper 1967), and can also be observed today directly above the

low sandy bluffs at the south ends of Monterey Bay and Point Reyes beaches (P.

Baye unpublished data 1997 to 1999). Today, little of the mobile sand perched

above the Merced bluffs is derived directly from the beach, as is indicated by the

lack of wind-shadow dune deposits on the bluffs. These perched dunes have

some soil characteristics of the much older, weathered sand deposits from which

they were derived. Similarly, the large climbing dunes at the north end of Baker

Beach appear to be relict deposits of a truncated, formerly more extensive dune

system, with low modern rates of sand transport directly from the beach. This

condition is indicated by landward-dipping beds in erosional dune scarps at the

foot of the dune, weathered mineral films on the dune sand, and the lack of windshadow sand deposits at the back of the beach (P. Baye unpublished data 1998). 

In contrast, Ocean Beach, along the western shore of the City, periodically

develops high rates of onshore wind transport of sand during phases of natural

shoreline widening (P. Baye, pers. observ. 1984 to 1999). Thus, some San

Francisco dune remnants today are truly relict (residual from former

environments, not geologically regenerated by modern physical and ecological

processes), while some dune remnants are actively geologically regenerated by

modern processes. 

The topography, stability, and soil of San Francisco dune remnants are also highly

modified by residual effects of past introduction of dune-stabilizing vegetation,

even where the original planted vegetation has vanished. Dune stabilization in

San Francisco began in the 1870's with the introduction of European beachgrass

(Ammophila arenaria), which successfully stabilized otherwise uncontrollable

mobile dunes and created steep, hummocky topography. Yellow lupine (Lupinus

arboreus), native to the region, was planted to enrich sandy soils with organic

nitrogen, in preparation for tree plantings. Iceplant (Carpobrotus edulis and

hybrids) strongly stabilizes both mobile and relatively stable dunes, and planted

stands of trees and shrubs (e.g., Monterey cypress [Cupressus macrocarpa], blue
1

 This reference is a citation of an herbarium specimen. Abbreviations for herbaria

are provided in the glossary, Appendix IV.

9

gum [Eucalyptus globulus], and wattle [Acacia spp.]) (U.S. Fish and Wildlife

Service 1998b, McLaren 1924) act as strong baffles to dune-forming winds, and

add abundant organic matter to dune soils.

b. Native Vegetation and Flora of the San Francisco Dune System. The original

San Francisco dune system supported a species-rich vegetation and contained a

wide spectrum of dune forms and successional stages. The character of the

historic dune system can be inferred approximately from: (1) modern

observations of comparable, protected dunes of the central coast; (2) historic

botanical records (Bolander 1863, Brandegee 1892, Kaufeldt 1954, Howell et al.

1958, and old herbarium collections); and (3) historic photographs (Greg Gaar

unpublished photographic archives 1999). Historic descriptive accounts and

interpretations (Ramaley 1918, Cooper 1967) are also valuable sources of

information. The plant species reported or collected from these dunes are shown

in Appendix I.

The outermost (western) parts of the system along Ocean Beach consisted of

mobile, mostly unvegetated sand masses (irregular wave-like transverse dunes)

and relatively sparse, hummocky vegetated foredunes of various dimensions, with

deflation plains (broad shore-parallel moist depressions) between them. Foredune

plants were probably limited to the few species capable of tolerating conditions of

strong sand mobility, such as beachbur (Ambrosia chamissonis), yellow sandverbena (Abronia latifolia), and occasionally dunegrass (Leymus mollis) or beach

saltbush (Atriplex leucophylla) at the seaward end. Silvery beach-pea (Lathyrus

littoralis) occurred at least locally near Lake Merced, probably in relatively

sheltered beach areas subject to occasional flooding. Pink sand-verbena (Abronia

umbellata, probably mostly introgressants with Abronia latifolia) was also present

in foredunes at least locally (collected by Peter Rubtzoff in 1959; CAS 5385981

),

as at Crissy Field, Presidio, today.

Dune slacks are topographically low, wide, and relatively flat areas within dune

systems, most of which are wetlands (Ranwell 1972). These important elements
10

of the dune ecosystem vary in their modes of origin and degree of wetness,

ranging from permanent ponds or dune lakes (e.g., Oso Flaco Lake, San Luis

Obispo County) to seasonally wet rush meadows. Many dune slacks originate as

deflation plains or hollows formed when wind erodes the dry sand surface down

to the moist capillary fringes of fluctuating water tables, or down to resistant

surfaces such as buried soil or lag deposits (concentrations of coarse sediment left

behind following erosion of finer particles) (Carter 1988, Ranwell 1972). 

Deflation slack complexes are best developed in Oregon (Wiedemann 1965), but

occur on the dunes of the central California coast at Dillon Beach in Marin

County, and Tenmile Dunes and Manchester Dunes in Mendocino County

(Cooper 1967, P. Baye unpublished data 1997). Dune slacks of the central

California coast are also often associated with springs and seasonal streams

perched on relatively impermeable sediment deposits beneath dune sheets, or

drainages from adjacent terrestrial surfaces or aquifers (Cooper 1967, P. Baye

unpublished data 1996 to 1999). Good examples of spring- or stream-fed dune

slacks analogous with those formerly in San Francisco occur within the central

coast region today at Point Reyes, Dillon Beach, Año Nuevo, and Franklin Point. 

Stream- or spring-fed slacks can be dammed by dunes, creating large seasonal or

permanent ponds, riparian willow/waxmyrtle thickets, or marshes. 

In San Francisco, there is historic evidence of both deflation and dammed-stream

types of slacks. Major ponded dune slacks occurred in the lee of major dune

ridges (Cooper 1967, Clary 1980), developing diverse rush/bulrush/sedge marsh,

ponds, and swampy thickets of willows (Salix spp., mostly arroyo willow,

S. lasiolepis) and waxmyrtle (Myrica californica). Good examples of dune slack

communities dominated by rushes, sedges, and forbs (herbaceous plants) are

found at Point Reyes, Dillon Beach, and Año Nuevo/Franklin Point. These

communities are probably like the ones formerly found in San Francisco. Dune

localities of many marsh, submerged aquatic, and wet-meadow plant species, and

descriptions of flats or hollows within dunes, were given by Brandegee (1892)

and Howell et al. (1958) (see Appendix I). The only dune slacks remaining in

San Francisco today are: (1) some of the ponds in Golden Gate Park (highly

altered derivatives of the original natural ponds of dammed dune slacks), and (2)

Mountain Lake in the Presidio.
11

Landward of the active transverse dunes, relatively more stable dunes occurred in

various stages of succession and rejuvenation (remobilization). They supported

either extensive stable cover of dune scrub or grassland vegetation, or a mosaic of

blowouts (hollows derived from wind erosion of dunes) within a matrix of dune

grassland and dune scrub. Dune scrub historically was, and still is, the dominant

vegetation of stabilized dunes. Dune scrub communities of the central California

coast have distinctive patterns of plant species abundance and composition that

distinguish them from coastal scrub of bluffs and hills lacking dune sand. 

Dominant woody (to subshrubby) dune scrub species in San Francisco included

false heather (Ericameria ericoides), silver or Chamisso’s dune lupine (Lupinus

chamissonis), coyote brush (Baccharis pilularis, a common coastal scrub

species), dune wormwood (Artemisia pycnocephala; probably in areas of

secondary blowouts), deerweed (Lotus scoparius), dune knotweed (Polygonum

paronychia), and dune buckwheat (Eriogonum latifolium) (Bolander 1863, Behr

1888, Brandegee 1892, Kaufeldt 1954, Howell et al. 1958). The yellow bush

lupine (Lupinus arboreus), which today is commonly found in dune scrub

remnants and restoration sites, was apparently not a component of the San

Francisco dune vegetation at the time of Bolander’s early surveys (Bolander

1863). It was formerly a minor component of central California coastal dune

scrub (Cooper 1936), although it was later broadcast-seeded into dunes to

stabilize them (McLaren 1924, Clary 1980). Other shrubs, such as poison-oak

(Toxicodendron diversilobum), California buckthorn (Rhamnus californica), and

lizard-tail (Eriophyllum staechadifolium, a species typical of coastal scrub in

general) were probably at least locally abundant elements of dune scrub; these

species also persist locally today in San Francisco dune remnants. Oak

woodlands dominated by dwarfed, shrubby coast live oaks also developed locally

in older dunes; very few persist today in some relict dunes of the Presidio (north

of the Public Health Services Hospital) and Golden Gate Park (around Fuchsia

Dell and De Laveaga Dell). Another dominant shrub of coastal headlands of the

Golden Gate, blue-blossom or California lilac (Ceanothus thyrsiflorus), is

conspicuously absent on dune substrates.

Common herbaceous species associated with San Francisco dune scrub include

man-root (Marah fabaceus), bracken fern (Pteridium aquilinum), live-forever

(Dudleya farinosa), miner’s-lettuce (Claytonia perfoliata ), maritime bromegrass
12

(Bromus carinatus var. maritimus), and many other species also found in dune

grassland (see Appendix I, and below). Indian paintbrush (Castilleja affinis ssp.

affinis) was formerly a common element of stable dune scrub in San Francisco

(Howell et al. 1958; then treated as Monterey Indian-paintbrush, Castilleja

latifolia), but today, in scrub remnants where it was formerly recorded, it is

infrequent and local (Howell et al. 1958, P. Baye unpublished data 1997 to 1999). 

A more common coastal Indian-paintbrush, Wight’s Indian-paintbrush (Castilleja

wightii), appears to be absent from San Francisco dunes (though local Indianpaintbrushes are sometimes easily confused with this strongly sticky-glandular

species), despite its frequency on San Mateo County coastal bluffs from Pacifica

south and its former occurrence near Lake Merced and Mt. Davidson (Howell et

al. 1958). Bare or sparsely vegetated gaps within coastal dune scrub intergrade

with dune grassland, and support annual herbs such as popcorn-flower

(Cryptantha leiocarpa), coast fiddleneck (Amsinckia spectabilis), wild heliotrope

(Phacelia distans), dune gilia (Gilia capitata ssp. chamissonis), San Francisco

spineflower (Chorizanthe cuspidata), annual plantain (Plantago erecta, usually

on older dune soils), and annual evening-primroses (Camissonia contorta and

Camissonia micrantha) (Howell et al. 1958; P. Baye, pers. observ. 1984 to 1999). 

Other species found in the dune scrub plant community in Presidio dune remnants

are the bush monkey flower (Mimulus aurantiacus), California bee plant

(Scrophularia californica), and toad flax (Linaria canadensis) (Brian O’Neill,

National Park Service, in litt. 2002).

Dune grassland and forbland intergrade with central coast dune scrub vegetation. 

Dune grassland and forbland are characterized by prevalence of grasses and

grasslike species or herbaceous plants (forbs) rather than the woody species of

dune scrub. Many of the same species occur in different relative abundance in

dune scrub. Stands of dune grassland and forbland are still well-represented in

other central coast dune systems (Point Reyes and Dillon Beach dunes, Monterey

Bay; P. Baye, pers. observ. 1984 to 1999), but are now poorly represented in San

Francisco. Small and recently-formed stands of dune grassland occur above

northeast Baker Beach, and in small remnant patches at Sunset Heights (P. Baye,

pers. observ. 1998). 
13

Among the common to dominant perennial grasses and grasslike plants of dune

grassland (also in openings in dune scrub) in San Francisco were dune bluegrass

(Poa douglasii), pacific wildrye (Leymus pacificus), red fescue (Festuca rubra),

maritime brome (Bromus carinatus var. maritimus), and at least locally, salt rush

(Juncus lesueurii; sometimes interpreted as Juncus breweri, both closely related

to wire rush, Juncus balticus) (Brandegee 1892, Howell et al. 1958). 

Herbaceous perennial plant species of dune grassland included dune goldenrod

(Solidago spathulata), sand mat (Cardionema ramossisimum), California poppy

(Eschscholzia californica—perennial dune ecological race), beach strawberry

(Fragaria chiloensis), California phacelia and common phacelia (Phacelia

californica, Phacelia distans), sea-pinks (Armeria maritima), yarrow (Achillea

millefolium), man-root or marah (Marah fabaceus), beach evening primrose

(Camissonia cheiranthifolia), as well as creeping perennial foredune species. The

tall, colony-forming dune-tansy (Tanacetum camphoratum) occurred in dune

scrub or forbland at the southern limit of its range in San Francisco. 

Common annual herbs included within dune grassland were essentially the same

as those in gaps within dune scrub, listed above and in Appendix I. Dune

grasslands also included species now rare or extirpated in local dune remnants,

such as the showy colony-forming broadleaf purple owl’s-clover (Castilleja

exserta ssp. latifolia; not recently reported), and sky lupine (Lupinus nanus; now

locally rare on dunes) (Brandegee 1892, Howell et al. 1958). 

Ecotones (transitional edges) of dune plant communities occurred where dune

sand deposits thinned out to veneers over bedrock, ancient Pleistocene sand

deposits, or dune slacks, with corresponding intermediate variations in plant

species associations. Remnant examples of dune ecotones in San Francisco are

found at Sunset Heights hilltops and around Sutro Heights (P. Baye, pers. observ.

1993 to 1999). The turf-forming sedge Carex praegracilis and the colonial rush

Juncus lesueurii and its allies probably occurred in dune/dune slack ecotones;

these species were present in San Francisco and occur in dune slacks elsewhere

on the central coast (P. Baye unpublished data 1995 to 1999).
14

Other types of sand deposits on the northern San Francisco Peninsula also support

many elements of the region’s coastal dune vegetation. Ancient, weakly

consolidated sandy deposits of sediment from “fossil” alluvial, lake, beach, dune,

and estuarine environments of the Merced and Colma formations (Schlocker

1974) date from the Pleistocene epoch. They are extensively exposed in San

Francisco and Daly City (Bonilla 1965, Schlocker 1974). Merced and Colma

deposits typically form yellow-brown coastal bluffs that erode and supply sand to

modern beaches and dunes, and directly merge with them in places (e.g., Fort

Funston, north end of Ocean Beach). They also occur at inland locations beyond

the Holocene (post-glacial) dune sheet (e.g., east of Lake Merced, Daly City, and

elsewhere; Bonilla 1965). The Colma sediments were probably derived from

extensive tidal lagoon sand shoals and adjacent barrier beach and dune

environments (Schlocker 1974). Some sands within these deposits resemble wellsorted dune sands, but contain significantly more clays, silts, and mineral

weathering products, giving them more soil-like properties. Coastal scrub and

grassland vegetation similar in composition and structure to those of coastal

dunes is associated with these Pleistocene sand deposits, including the

southernmost locality of Lessingia germanorum (McClintock et al. 1990).

c. Dynamics of Dune Vegetation. The structure and composition of dune scrub

and grassland vegetation are closely connected to the geomorphic dynamics of

sand transport within dune systems, particularly in relation to dune blowouts. 

Blowouts in dunes that have previously been stabilized by vegetation and initial

soil development reactivate mobility of buried sand, creating zones of upwind

erosion and downwind sand accretion. Blowouts establish patches of secondary

vegetation succession, gaps in which local pioneer vegetation can establish. 

Blowouts arise through various interacting influences, such as animal burrowing,

fire, and drought-related dieback of vegetation. They occur at various scales,

ranging from minor patches a few meters wide, to reactivation of whole ridges

(Cooper 1958, Ranwell 1972). Blowouts of different sizes and ages within a

matrix of dune vegetation undergo secondary succession from dune scrub back to

dune forbland or grassland. The variation in size and ecological maturity of

blowouts within dune systems can maintain a mosaic of vegetation gaps available

for recolonization, and diverse vegetation types arising from variable local

influences and stages of development. Most blowouts in San Francisco dune
15

remnants appear to be the result of recent human disturbances, especially

trampling of vegetation (P. Baye, pers. observ. 1984 to 1999).

Blowouts in modern dunes of the central California coast are impeded by dense

cover of invasive sand-stabilizing vegetation, particularly European beachgrass

(Ammophila arenaria), deliberately introduced to San Francisco around 1870 for

dune stabilization purposes; Jepson 1911, McLaren 1924, Clary 1980, U.S. Fish

and Wildlife Service 1998b) and iceplant (Carpobrotus edulis) and related

hybrids. By the mid-20th century, these and other nonnative species dominated

remnant San Francisco dune vegetation (Kaufeldt 1954). Blowout vegetation

development is also thwarted by wind-sheltering effects of large nonnative trees

(particularly Monterey cypress [Cupressus macrocarpa], and blue gum

[Eucalyptus globulus], planted on San Francisco dunes for windbreaks and

planted “forest” cover; McLaren 1924, Clary 1980). The elimination of

burrowing, grazing, and trampling activities by native mammals also probably

contributed to the reduction of blowouts in remnant old dunes in San Francisco. 

In contrast, intensive recreational use of dunes, particularly in foredune areas near

beaches, can either compensate for the over-stabilizing effects of nonnative

vegetation, or over-compensate and exaggerate the distribution and abundance of

mobile, unvegetated sand. At Ocean Beach and Fort Funston, intensive trampling

and footpaths frequently enlarge to blowouts. Since the construction of the Great

Highway along Ocean Beach in the foredune zone, the foredunes have been

periodically stabilized by placement of earthen fill material and plantings of

European beachgrass over most of its length.

2. San Franciscan Bedrock Outcrops and their Vegetation

a. San Francisco Bedrock Outcrops. Bedrock outcrops occur scattered

throughout San Francisco (Schlocker 1974). Many persist as steep,

undevelopable knobs on the crests of hills up to 281 meters (922 feet) above sea

level, or high, unstable coastal bluffs subject to frequent landslides (Schlocker

1974). The bedrock outcrops in San Francisco are folded and uplifted Jurassic

and Cretaceous sedimentary rocks with intrusions of igneous (volcanic) rocks of

the Franciscan formation. Franciscan rocks include massive and bedded

sandstones, shale, chert, greenstone (mostly basalts), serpentinite, gabbro-diabase,
16

and mixed sheared rocks along fault zones. They range from erosion-resistant

basalt and chert, to serpentine rocks that range from massive, hard, and dense to

intensely sheared, soft, friable, and plastic material (Schlocker 1974). Serpentine

rocks are derived from weathering or modification of igneous ultramafic rocks

(rocks originating as deep magmas rich in heavy metals). Serpentine rocks are

particularly relevant to distinct local vegetation, since the soil chemistry of

serpentine typically imposes extremes of mineral nutrition and toxicity

(particularly low calcium, high magnesium, and heavy metals) for many plants. 

Serpentine soil results in exclusion or growth suppression of many plant species

and selects for a narrow range of endemic serpentine-adapted species or

populations. Many serpentine plants are narrow edaphic endemics, species of

extremely restricted ecological and geographic distribution based on soil type

(Kruckeberg 1954, 1984; Raven and Axelrod 1978; U.S. Fish and Wildlife

Service 1998a). 

Most bedrock outcrops of the interior parts of San Francisco are characterized by

steep topography, thin, dry soils, and bare rock, conditions that maintain 

permanently sparse vegetative cover, at least locally. The extensive serpentine

soils and rocks of the Presidio bluffs, along the south shore of the Golden Gate

(Figure 2 and Figure 3) occur in both structural outcrops and in landslides. At

some locations in the bluffs, the exposed serpentine has deeper soft soils that

undergo extremes of saturation and drying, frequent local seeps and near-surface

groundwater, as well as stronger direct exposure to marine fog (P. Baye, pers.

observ. 1984 to 1999). The largest outcrops of serpentine occur on Potrero Hill

(Schlocker 1974). Major outcrops of sheared rocks and greenstone are shown in

Figure 3. Significant bedrock outcrops occur at Mount Davidson Park, Twin

Peaks, Mount Sutro, the Duboce U.S. Mint, Sunset Heights, McLaren Park, Point

Lobos, and other locations (Figure 2 and Figure 3), many of which are partially

obscured by nonnative vegetation, or are isolated as small erratic remnants

included within developed areas (e.g., serpentine at the Duboce U.S. Mint and

near China Basin). Significant historic outcrops, now destroyed, occurred at the

Masonic and Laurel Hill cemeteries in the Richmond district and at the former

Protestant Orphan Asylum at Haight and Laguna streets, near the Duboce U.S.

Mint (U.S. Fish and Wildlife Service 1984).
17

Figure 2. Localities of serpentine and greenstone bedrock outcrops in San

Francisco. Areas shown include urbanized and undeveloped sites. 

Some otherwise urbanized sites include local outcrops (road cuts,

graded slopes) that support vegetation. Approximate localities of

historic Raven’s manzanita sites are indicated by circled stars (k).
18

Figure 3. Geological control of Raven’s manzanita habitat. Distribution of

shear zones, containing sheared rocks with serpentine outcrops, and

Franciscan formation outcrops including greenstone. Reproduced

from Schlocker (1974) (Figure 63). 
19

b. Vegetation of San Francisco Bedrock Outcrops. The vegetation associated

with bedrock outcrops in San Francisco is variable today, including elements of

remnant native vegetation as well as naturalized nonnative vegetation. 

Historically it included plant associations classified as coastal grassland (prairie)

and variations of coastal scrub (Appendix II). No native trees are known to have

originally been associated with serpentine or other bedrock outcrops in San

Francisco (Bolander 1863, Cooper 1875, Behr 1892, Brandegee 1892), although

introduced Monterey cypress, Monterey pine (Pinus radiata), and blue gum have

colonized serpentine landslides of the Presidio (P. Baye, pers. observ. 1998). 

Many nonnative herbaceous species are able to colonize bedrock outcrops,

including serpentine rocks. Species that are particularly invasive locally include

jubata grass (Cortaderia jubata), filaree (Erodium spp.), iceplant (Carpobrotus

edulis and hybrids), wild oat (Avena fatua), acacia species (Acacia species), and

Cape ivy (Delairia odorata) (U.S. Fish and Wildlife Service 1984; P. Baye, pers.

observ. 1993 to 1999; and Brian O’Neill, National Park Service, in litt. 2002). 

Invasive species that are able to colonize outcrops abundantly can eliminate the

characteristic open, barren, or sparse structure of the vegetation. Slow-growing,

short-stature native species, including serpentine endemics, probably are able to

persist because of this characteristic open, low vegetation structure (Kruckeberg

1984), and are threatened by competition and habitat modification due to invasive

nonnative plants (U.S. Fish and Wildlife Service 1984, 1998a).

In San Francisco, no unique native plant assemblages are associated with bedrock

outcrops, even those on serpentine. However, many species and local variations

in plant associations have distributions closely corresponding to these habitats

(U.S. Fish and Wildlife Service 1998a). The historically documented distribution

of local manzanita species was closely correlated with relatively barren bedrock

outcrops, particularly of serpentine and greenstone (U.S. Fish and Wildlife

Service 1984, Figure 2). Similarly, Bolander (1863) noted that some other

manzanita species were closely associated with sandstone outcrops of the

Oakland Hills, and Howell (1949) observed that most manzanita species in Marin

County were associated with rocky, barren, steep slopes and ridges. Some thistles

are frequently associated (Franciscan thistle, Cirsium andrewsii) or exclusively

associated (Mount Tamalpais thistle, Cirsium hydrophilum var. vaseyi, and

fountain thistle, Cirsium fontinale var. fontinale) with serpentine seeps (Keil and
20

Turner 1993). Rare or uncommon species in the San Francisco flora that are

relatively frequent on or exclusive to serpentine or other bedrock outcrops include

Marin dwarf-flax (Hesperolinon congestum), evax (Hesperevax sparsiflora), and

San Francisco gumplant (Grindelia hirsutula var. maritima) (Howell et al. 1958,

Golden Gate National Recreation Area unpublished reports). Many other

grassland species that formerly occurred in greater abundance in San Francisco

(Wood 1996) are now restricted to the thin soils and steep slopes that have

marginally escaped the adverse influences of urban development and nonnative

vegetation.
21

II. BIOLOGY OF FEDERALLY LISTED SPECIES

A. San Francisco Lessingia (Lessingia germanorum Chamisso)

1. Description and Taxonomy

San Francisco lessingia is an annual herb in the aster family (Asteraceae) (Figure

4). Seedlings and young vegetative plants (Figure 4a) develop from unbranched

rosettes of oblanceolate (spear-shaped but tapered at base, wide and rounded at

tip) leaves. As the plant matures (Figure 4b), some lateral shoots elongate

markedly and then branch profusely, developing a decumbent (growing mostly

through lateral branches) growth habit. Mature plant heights can range from less

than 5 centimeters (2 inches) in stunted plants, to 0.3 meter (1.2 feet) tall. Mature

stems are reddish brown and are tomentose (with loose grayish woolly hairs). 

Leaves on mature stems are small (0.5 to 3.0 centimeters [0.2 to 1.2 inches]),

most less than 1 centimeter (0.3 inch), pinnately lobed (branching from a single

central vein), toothed or entire (lacking teeth and lobes), oblanceolate or longtapered obovate (egg-shaped, but widest at the far end), and grayish-green due to

dense woolly hairs (Figure 4c). Flowers in the aster family occur in flowerheads

composed of many individual flowers (florets). Flowerheads function as

individual units for pollinators and resemble individual flowers. In San Francisco

lessingia, flowerheads appear from late summer through fall. They occur singly

or in loose clusters at the ends of stems (Figure 4d). Depending on plant size,

individuals may bear a few to hundreds of flowerheads. The bell-shaped

involucre (a mantle of tiny leaf-like appendages called phyllaries that enclose the

immature flowerhead) is 4 to 8 millimeters (0.15 to 0.3 inch) wide. Phyllaries are

lance-shaped, with abruptly pointed tips. All florets within the flowerhead are

disk florets (composite flowers within the flowerhead); ray florets (small

individual flowers with strap-shaped corollas resembling single petals at the

margin of the flowerhead) are lacking (Figure 4e). Each head contains from 20 to

40 disk florets. Each floret has a yellow corolla (series of petals that are united

into a tubular to funnel-shaped deeply lobed structure), with brownish bands in

the throat. The fruit is an achene (a seed-like dry fruit) 1 to 3 millimeters (0.04 to

0.12 inch) with tan or whitish pappus (hairy bristles that increase the 
22

Figure 4. San Francisco lessingia (Lessingia germanorum). (a) unbranched

juvenile plant; (b and c) branching juvenile plant, pre-flowering stage; (d) detail,

vegetative shoot; (e) detail, flowerhead; and (f) mature plant, early flowering

stage.

f

c

d

a b

e
23

dispersability of achenes in air currents, as in dandelion “blowballs”). No other

annual Lessingia species occur within the range of San Francisco lessingia on the

San Francisco Peninsula, making it easy to distinguish (Howell 1929, Spence

1964, Lane 1993). San Francisco lessingia was originally collected and described

as a species by Chamisso in 1829, based on specimens collected from sand dunes

in San Francisco (Howell 1929), probably from the Presidio. Lessingia

germanorum is the type species for its genus (the first plant selected by

taxonomists to represent the group; the exemplar for all related species). Howell

(1929) treated San Francisco lessingia broadly as a taxonomic entity, assigning all

of the yellow-flowered taxa of Lessingia to 11 varieties of Lessingia

germanorum. Howell treated the endemic San Francisco lessingia taxonomically

as Lessingia germanorum Cham. var. germanorum. Meredith A. Lane’s

treatment of the genus Lessingia in California for the Jepson Manual (Hickman

1993) returned San Francisco lessingia to a narrow species concept, eliminating

Howell’s varieties by assigning them to other Lessingia species or elevating them

to species rank. San Francisco lessingia is distinguished from other California

species of Lessingia by the combined presence of yellow disk florets and lack of

ray florets, and the general absence of glands on the foliage (or having only

sparse, nail-shaped glands). Howell (1929) considered San Francisco lessingia to

be a slightly modified form of the most primitive (ancestral) types of Lessingia.

San Francisco lessingia is currently considered a distinct species, it was formerly

thought to be most closely related to Lessingia glandulifera A. Gray var.

pectinata Jeps. (Howell 1929, S. Markos, pers. comm. 1998). Hoover (1970)

interpreted Eastwood specimens of Lessingia from the Santa Maria River bed in

San Luis Obispo County as Lessingia germanorum var. germanorum (following

Howell’s classification), based on the nearly glandless condition. This specimen

was collected within the range of Lessingia glandulifera var. pectinata (treated by

Hoover as Lessingia germanorum var. pectinata (E. Greene) J. Howell), which

occurs in essentially the same type of open, sandy coastal habitat, but in a

restricted southern range (Howell 1929). The latest genetic analysis indicates that

San Francisco lessingia is a member of a clade (a group of species descended

from a common ancestor) comprised of Lessingia tenuis, Lessingia glandulifera

var. pectinata, and some members of Lessingia germanorum, with the resolution

of its nearest relative within that group requiring further work. Reevaluation of
24

the genus based on further molecular genetic data is likely to clarify systematic

relationships within Lessingia, particularly with Lessingia glandulifera var.

pectinata (Staci Markos, Jepson Herbarium, U.C. Berkeley, in litt., 2002; and

Markos and Baldwin 2002).

2. Historic and Current Distribution and Abundance

In the 19th century, San Francisco lessingia was reported or collected from

numerous unspecified localities and two specific areas in San Francisco: one in

the northwest, from the Presidio (near Lobos Creek) to Lone Mountain, and one

in the southwest, near Lake Merced (Brandegee 1892, Table 1, Figure 1). Most

early collection localities were highly generalized as “Presidio” or “San

Francisco.” Herbarium sheets with specific collection localities from the 19th and

20th centuries clearly indicate that the historic distribution of San Francisco

lessingia on the San Francisco Peninsula was considerably wider than today

(Table 1). The modern restricted distribution of San Francisco lessingia is

probably due to habitat loss, habitat alteration, and extirpation of populations

(Spence 1964). 

The northwestern San Francisco population of San Francisco lessingia included

specific collection localities from dunes as far east as Fulton Street at Lone

Mountain (J. T. Howell [1926], CAS 166038, [1927] CAS 166064; earliest

collections here in the 1860's: J.P. Moore, no date, circa 1860's, CAS 9451; A.

Kellogg & W. G. W. Harford [1868], CAS 9448) and as far west as Baker Beach

at the mouth of Lobos Creek (CAS 491875, P.H. Raven [1954]). Some of the

historic Presidio localities appear to be the same as those found today (Spence

[1960] “sandy soil, roadside adjacent to golf links,” CAS 567670; “sandy area

between U.S. Public Health Services Hospital and Lobos Creek,” CAS 188634, P.

Rubtzoff [1956]), but some are apparently extirpated (“Mountain Lake,” CAS

128600, Goodman [1954]); “Coastal bluffs near mouth of Lobos Creek,” (Raven

[1954], CAS 491875) (Table 1).

a. Modern Presidio Populations. The Presidio populations are currently located

at six sites, each one a few tens to hundreds of meters from the nearest

neighboring colony (Figure 5). Population estimates from the Presidio are 
25

Table 1. Historic locality and habitat data 

from herbarium specimens of Lessingia germanorum

Where duplicated specimens exist, label information from the more detailed account is cited. Bold

type is added, not in original.

CAS - California Academy of Sciences

DS - Dudley/Stanford (at CAS)

JEPS - Jepson Herbarium (at UC Berkeley)

UC - University of California (Berkeley)

______________________________________________________________________________

L. Abrams, Aug 31, 1901. CAS 145061. Sand dunes, San Francisco County.

H.G. Bloomer, 1868. CAS 190998. San Francisco.

T.S. Brandegee, July 1, 1890. UC 87998. Marine Hospital, San Francisco.

K. Brandegee, Sept. 1, 1901. UC87997. San Francisco.

K. Brandegee, July 12, 1905. UC 83863. Lake Merced, upper end. San Francisco.

K. Brandegee [as “MKC”], Aug. 1, 1886. JEPS16551. San Francisco.

K. Brandegee (no date) CAS 6438. San Francisco.

E. Cannon (no date) CAS 9452. Presidio Grounds (San Francisco).

J.W. Congdon, May 1881. CAS 6436. San Francisco.

W.R. Dudley, Nov 8, 1899. DS 593657. Ocean View Downs, San Francisco.

A. Eastwood, Aug. 7, 1913. CAS 3449. Lake Merced, San Francisco.

Goodman, Dr [sic]. Dec. 1, 1924. CAS 128600. Mountain Lake, San Francisco.

J.T. Howell, Oct. 19, 1924. CAS 166017. near Lake Merced, San Francisco.

J.T. Howell, Aug. 12, 1926. CAS 166021, UC 912539. Near Lone Mountain, in deep sand.

J.T. Howell 2233, Nov. 7, 1926. CAS 166038, JEPS 20555. Sandy open [....] near Lake Merced,

San Francisco.
26

Table 1, continued.

J.T. Howell, Mar. 18, 1927. CAS 166064. Fulton Street east of St. Ignatius Church, San

Francisco, a part of Lone Mountain slopes.

J.T. Howell, Sept. 10, 1927. CAS 166053. Grown in the Botany Garden, University of California,

in Strawberry Cañon, from seed collected at Lake Merced, San Francisco. 

J.T. Howell, Sept. 26, 1931. UC 473579. Sandy slope, Presidio, San Francisco.

J.T. Howell 8126, Sept 29, 1931. CAS 188634. Sandy slope, Presidio. San Francisco.

Jepson, W.L. Oct. 1, 1894. JEPS 20553. Hills near Mountain View Lake [Mountain Lake, San

Francisco].

A. Kellogg & W. G. W. Harford, July 7, 1868. CAS 9448. Lone Mountain, San Francisco.

J. P. Moore, (no date, ca. late 1800's). CAS 9451. Lone Mt.

N. Pettibone, July 1894. CAS 346025. Presidio, San Francisco.

P.H. Raven, Aug. 22, 1954. CAS 491875, JEPS 16551. TOPOTYPE. Coastal bluffs near mouth

of Lobos Creek, Presidio, San Francisco.

L. Rose, Sept. 9, 1941. CAS 305722, UC662638. Type locality. Associated with Croton

californicus and Baccharis pilularis on coastal sand hills, Presidio, San Francisco. alt. 60 m.

L. Rose Oct. 21, 1961. DS 567812, JEPS 30082. Presidio, sandy flat. ca. 200 ft. [San Francisco].

L. Rose,. July 14, 1969. JEPS 59952. Presidio, n of golf links. Dry sandy flats.

P. Rubtzoff,. Aug 18, 1956. CAS 494874. Presidio Waste, sandy area between U.S. Marine

Hospital and Lobos Creek.

W. Spence, 12 June 1960. CAS 567670, UC 1287362. Flowers deep lemon yellow with a reddish

brown band in the throat. Growing in sandy soil, roadside adjacent to golf links, Presidio, San

Francisco.

Mrs. E. C. Sutcliffe, May 1918. CAS 9450. Presidio, San Francisco.
27

Table 1, continued.

______________________________________________________________________________

Summary of collection localities:

unspecified San Francisco localities 6

unspecified Presidio localities 7

Mountain Lake (Presidio) 2

Presidio Golf Links vicinity 2

Marine Hospital - upper Lobos Creek vicinity (Presidio) 2

Baker Beach, mouth of Lobos Creek (Presidio) 1

Lone Mountain 4

Lake Merced vicinity 5

Ocean View vicinity (Lake Merced?) 1
28

Figure 5. Approximate contemporary locations of San Francisco lessingia

(Lessingia germanorum) in the Presidio, San Francisco. Circled

stars (k) indicate approximate locations of remnant San Francisco

lessingia colonies. Outer circles indicate vicinity of probable or

actual population spread. 
29

provided from Golden Gate National Recreation Area file information. 

Population sampling and estimation methods used by Presidio Natural Resources

staff are described and evaluated by Wayne (1996) and Bode (1998). Site

locations are shown in Figure 5.

(1) The Lobos (Creek) Dunes site is an area of early-succession stable

dune scrub that was recently (1995 to 1997) restored as a mitigation effort

following the construction of the Richmond Transport Project. 

Restoration involved the removal of a ball field, a road, nonnative trees,

iceplant, and other herbaceous nonnative plants; the importation of sand

material; and replanting locally propagated native vegetation (J. Cannon,

M. Albert, pers. comm. 1996 to 1997; Brian O’Neill, National Park

Service, in litt. 2002; Craig Middleton in litt. 2002). This restoration

expanded the small (less than 0.2 hectare [0.5 acre]) remnant Lobos Creek

population that occurred on a formerly graded, disturbed, weedy, open

space on mixed dune sand and fill substrates above Lobos Creek. The site

is bordered by nonnative Monterey cypress groves and Lobos Creek’s

riparian zone. The restored dune area supporting San Francisco lessingia

now occupies approximately 5.3 hectares (13 acres). After regrading the

site and seeding with San Francisco lessingia, the population here has

recently expanded from a few tens to hundreds of plants in the 1980's to

an estimated 130,000 in 1997 and approximately 1,500,000 in 1998

(National Park Service 2003). As replanted native dune scrub vegetation

established after 1998, lessingia numbers adjusted to a lower, more

sustainable range between 340,000 and 1,500,000 lessingia plants

(National Park Service 2003). The rapid population changes of San

Francisco lessingia at this site reflects the early successional development

of the restored vegetation there.

(2) The Battery Caulfield Road site is a small dune scrub patch (0.4

hectare [1 acre]) that was subjected to early (1988) restoration efforts by

the Golden Gate National Recreation Area (National Park Service). A

planted Monterey cypress grove partially shades the site to the west and

isolates it from Lobos Dunes. The site is heavily invaded by nonnative

grasses. It supports a moderate sized colony of San Francisco lessingia,

consisting mostly of small plants (P. Baye, pers. observ. 1996 to 1999,
30

Golden Gate National Recreation Area unpublished data). Population size

has been estimated up to 4,800 plants in 1998, increased under

management from 873 in 1997. During the 1990's, population size ranged

between approximately 1,000 and 6,000 plants (National Park Service

2003), but declined to 824 plants in 2000 (M. Albert, unpublished data

2001). This population is reintroduced, the product of seeding conducted

in 1988 at the approximate location of a historic remnant colony (Golden

Gate National Recreation Area unpublished data). The original colony

was probably a subpopulation or daughter colony of the population at the

adjacent Lobos Creek site, from which it was separated by the Monterey

cypress grove that today covers many hectares of a high, west-facing

intact dune slope.

(3) The Public Health Services Hospital site, behind (northeast of) the

Public Health Services Hospital (which is located on the site of the former

Marine Hospital), is a disturbed remnant stable dune with coast live oak

(Quercus agrifolia), native and nonnative grassland, and dune scrub

vegetation (mostly mock-heather, Ericameria ericoides, and coyote-brush,

Baccharis pilularis). Part of this site had been disturbed by earthmoving

activities (a borrow pit for sand and haul road) and trampling, which may

have provided or expanded open sandy habitat for San Francisco lessingia

here in the past. The site is bordered by a strip of Monterey pine (Pinus

radiata) to the south and a strip of blue gum trees (Eucalyptus globulus) to

the east. These tree plantings function as wind-breaks and visual barriers

for the adjacent Presidio golf course. San Francisco lessingia is most

often associated here with partially bare erosional slopes, remnants of

vehicle and pedestrian tracks, annual-dominated sandy flats, and edges of

active pedestrian paths along fences — all microsites where vegetation is

either low, open, or sparse. The site, now fenced and relatively

undisturbed, shows indications of succession toward closed perennial and

woody native dune scrub vegetation, as well as colonization by some oak

seedlings (P. Baye unpublished data 1998). Recent estimated population

sizes (1990's to 2000) usually ranged from 3,000 to 19,000, but surged up

to 78,000 in 1998 after intensive weeding (National Park Service 2003)

occupying a variable area around 0.6 hectare (about 1.5 acres). San

Francisco spineflower (Chorizanthe cuspidata) and dune gilia (Gilia
31

capitata ssp. chamissonis), among other associates of San Francisco

lessingia, have occurred at the site since 1994. An undisturbed grove of

low-growing native oak woodland and dune scrub borders the west end of

the San Francisco lessingia population. A formerly

excavated/graded/filled dune area supporting mixed wet meadow and

dune slack vegetation (salt rush, Juncus lesueurii; Pacific blackberry,

Rubus ursinus), dune scrub (mostly coyote-brush), and iceplant

(Carpobrotus edulis) occurs to the north and northwest.

(4) The Presidio Golf Course roadside site is a narrow, steep road cut in

old dunes on the west side of the maintenance road adjacent to the

Presidio Golf Course, across the golf course from the Public Health

Services Hospital lessingia site. This area, which covers less than 0.4

hectare (1 acre), is a gap in the narrow stand of blue gum trees that

otherwise line the top of the cut and border the golf course. The steep

slope undergoes small-scale chronic erosion, and supports a moderate to

small population of San Francisco lessingia in the vegetation gaps

(ranging from 215 to 8,000 individuals, usually fewer than 2,000; Golden

Gate National Recreation Area unpublished data 1998 to 2001). 

Dominant vegetation comprises typical native dune scrub dominants,

nonnative grasses (primarily Briza sp.), and bracken fern (Pteridium

aquilinum) (P. Baye unpublished data 1998). The site is bounded by the

maintenance road, the Golf Course, and blue gum stands, and is parallel to

Highway 1. This population is located near Mountain Lake, an historic

extirpated locality of San Francisco lessingia. 

(5) The Rob Hill site is a patch of disturbed, weedy old dune scrub and

grassland at the edge of a blue gum grove, and adjacent to Battery

McKinnon-Stotsenberg. It supports a population of San Francisco

lessingia ranging in the thousands of plants. The portion of the site

supporting locally abundant dune annuals and San Francisco lessingia

covers about 0.4 hectare (1 acre). The site is bounded by a dense grove of

blue gums, an evergreen understory of Canary Island ivy (Hedera

canariensis), and historic military buildings. Population size ranged

around 3,000 to 9,000 individuals in the 1990's, but increased to more than

155,000 in 1998 under intensive management (primarily weeding of
32

nonnative annual grasses), and declined to fewer than 500 in 2000.

(Golden Gate National Recreation Area unpublished data 1998, M. Albert

and P. Holloran, pers. comm. 1998 to 2001). 

(6) The Wherry Dunes restoration site (referred to by the National Park

Service as the Feral Dunes), consists of about 1 hectare (2.5 acres) of

restored habitat, together with an area of dune scrub remnant referred to

by the National Park Service as “Feral Scrub”. This site currently

comprises 4 hectares (10 acres) above the north end of Baker Beach near

Battery Chamberlain, located upslope of Lincoln Boulevard at the north

end of Pershing Drive. The part of the site added to the habitat by

removal of demolished buildings has been planted with native dune scrub

vegetation, and a small, volatile (subject to large, rapid fluctuations in

size) founder population of San Francisco lessingia established

spontaneously, beginning with a few plants in 1997. It is believed that

San Francisco lessingia seed may have been dispersed on footwear and

clothing of numerous volunteer restorationists working at this site after

coming from working at the large, dense population at Lobos dunes (J.

Cannon, pers. comm. 1998), but it also is possible that it may have

dispersed without human assistance. The population expanded to several

thousand plants spread over a hectare (2.5 acres) in 1999 (P. Baye, pers.

observ. 1999) and nearly 150,000 plants in 2000 (National Park Service

2003).

Overall annual population size of San Francisco lessingia in the Presidio

increased progressively from approximately 21,000 individuals in 1994 to

a peak size of over 1.7 million in 1998 (National Park Service 2003). 

Most of this population change was due to the artificial seeding and

habitat expansion of the Lobos dunes restoration in an early successional

stage. The estimated population size has remained at less than one third of

the 1998 peak size since then. Other important factors coinciding with the

population increase were intensive vegetation management (manual

weeding, particularly at Rob Hill) and above-normal rainfall (other than

the early dry spring of 1997). These factors need to be considered during

implementation of the long-term strategy to recover San Francisco,

particularly when evaluating population numbers in relation to recovery

criteria.
33

b. Southwestern San Francisco and Vicinity Populations. The historic

southwestern San Francisco population included collection localities from the

Lake Merced area, probably including areas now within the Fort Funston dune

area immediately west of Lake Merced (CAS 3449, Eastwood 1913; CAS

166017, J. T. Howell 1924; CAS 66038, J. T. Howell 1926; Table 1). Collections

also were made at Ocean View (DS 593657; Dudley, 1899) southeast of Lake

Merced where Colma Formation sands occur east of the San Francisco dune sheet

(Figure 1). It is not clear whether the “Lake Merced” records were on the east

shore area (on Colma Formation sands outside the San Francisco dune sheet), or

on dune sands now associated with the modern Fort Funston dunes along the west

shore of Lake Merced. No records of San Francisco lessingia are known from the

eastern dunes of the city, nor from the extensive mobile dunes of the Sunset

district, which persisted well into the 20th century. The Lake Merced and Ocean

View populations are now extirpated, last collected there in the late 1920's.

(1) Hillside Park, Daly City (San Bruno Mountain) population. The

southwestern population of San Francisco lessingia is now represented

only by the Daly City population, first reported in 1989 by Elizabeth

McClintock (Golden Gate National Recreation Area unpublished files). 

This population is exceptional geographically and ecologically. It occurs

on the western end of San Bruno Mountain on slopes above Hillside Park,

below the south face of Reservoir Hill. This location is far south of the

main San Francisco dune sheet mapped by Cooper (1967) and Bonilla

(1965) (Figure 1 and Figure 6). The sandy substrate is derived from the

weakly consolidated and weathered Pleistocene sand deposits of the

Colma Formation, a composite of uplifted sands originally deposited in

ancient lagoon or estuarine environments. They are weathered and

strongly iron-stained (bright yellowish or rusty brown), and include some

silt and clay layers (Bonilla 1965, Schlocker 1974). 

The origin of the Hillside Park, Daly City population is uncertain. This

population may be derived from a local relict population, or it may

represent opportunistic modern colonization of regraded sandy slopes,

possibly from the former Ocean View-Lake Merced population of

southwestern San Francisco. The antiquity of this isolated population is

doubtful for several reasons. San Bruno Mountain had been botanized 
34

Figure 6. Approximate contemporary location of San Francisco lessingia

(Lessingia germanorum) at Hillside Park, Daly City. Circled star ( k) indicates

approximate location of San Francisco lessingia colonies.
35

intensively in the early to mid 20th century, but no one reported San

Francisco lessingia until recently from an area northwest of San Bruno

Mountain, other than the Ocean View collection. Also, the entire

population occurs on graded, terraced slopes. The species’s center of

abundance at the site is on recently excavated and redeposited sand along

a utility line, surrounded by older, dense coastal scrub that is unsuitable

habitat. However, whether it is a surviving relict population that persisted

at the Hillside Park location, or a more recent recolonization from the

Ocean View-Lake Merced population, the Daly City population, based on

recent genetic work, is distinct from those at the Presidio. The difference

was as great as the difference between the San Francisco lessingia at the

Presidio and some other species of lessingia (Markos and Baldwin 2002;

and Staci Markos, Jepson Herbarium, U.C. Berkeley, in litt., 2002).

Some of the graded slopes cut in the Colma Formation sands above

Hillside Park have subsequently been naturally reworked by wind into

dune deposits. The sand here is similar to that of the Fort Funston bluff

sediments (mostly Merced formation), which supply much of the parent

material for the secondary dunes perched on top of them. These old ironstained yellowish sands contrast with the gray to tan recent Holocene

sands of the San Francisco dune sheet. The Hillside Park sand slopes

support coastal scrub vegetation that is essentially similar to the dune

scrub of San Francisco, particularly the Sunset Heights dune remnants

(Hawk Hill, Sunset Heights Park, Grandview Park) and Fort Funston

dunes (P. Baye unpublished data 1998).

The Hillside Park slope population of San Francisco lessingia extends

from the top of the east-facing slope at the end of Bismarck Street to the

end of the south-facing slope near Price Street and the Kennedy School,

above Bonnie Street (Figure 6). It is the largest of the “natural” (not

deliberately restored or seeded) remnant populations in terms of

population size and area, exceeding even the Rob Hill site. The highest

densities of San Francisco lessingia in 1999 occurred in the vicinity of a

seep and willow grove near a buried utility line alignment leading to the

crest of Reservoir Hill from the corner of Bonnie and Lausanne streets. 

The roughly linear high-density strip of San Francisco lessingia and dune
36

annuals probably tracks the footprint of past grading activities along the

buried pipeline alignment. Low densities (scattered individuals and small

colonies) occur near the summit of a weedy east-facing slope near blue

gums, iceplant, and residential back yards on Wyandotte Avenue (as far

south as Bismarck Street); and on the graded, terraced slope above Bonnie

Street (P. Baye unpublished data 1998). The population occurs on both

private land and municipal lands of Daly City. Portions of the site have

been maintained in recent years by citizen volunteers using manual

removal methods to control invasive nonnative vegetation (S. Smith, pers.

comm. 1998).

c. Variation in Abundance of San Francisco Lessingia. Population sizes of the

various San Francisco lessingia sites have fluctuated extremely since they have

been observed during the last two decades, apparently in response to factors such

as drought, mechanical disturbance, artificial seeding (augmentation and

reintroduction), weeding activities, and competition with weeds. In 1980, the

single population then known (now within the Lobos Dune site) included only

250 plants. The population decreased to 19 plants in 1985, the most extreme

population “bottleneck” since monitoring began. Population size rose to around

900 in 1986, and declined to approximately 600 in 1989 (Golden Gate National

Recreation Area file information). In 1994, after years of improved weed

management efforts and discovery of additional local populations, the cumulative

Presidio population approached an estimated 20,000 plants. In 1998, after 2 years

of restoration at Lobos Creek Dune site (involving extensive tree removal,

grading, and direct seeding of disturbed sand) and intensive weeding at the other

population sites, overall population size of San Francisco lessingia in the Presidio

(total of all sites) peaked at roughly 1.7 million; the current population size as of

2001 is under 0.5 million (National Park Service 2003). In 1989, only 310 plants

were known from the Hillside Park site. In 1998, the population above Hillside

Park was at least an order of magnitude larger than the 1989 estimate (P. Baye

unpublished data 1998). The overall rangewide number of San Francisco lessingia

plants, though highly fluctuating and quantified with variable accuracy and

precision, ranges over a magnitude around 500,000 to 2,000,000 plants. 

Population size and population trends are sensitive to habitat conditions,

particularly the availability of sparsely vegetated to bare sand within dune scrub
37

and grassland. The species is distributed over seven relatively discrete

populations, with the largest (core) population at the restored Lobos Dune site.

3. Ecology and Reproduction

a. Environmental Conditions. Modern populations of San Francisco lessingia

have high fidelity to vegetation gaps in stabilized old sand dunes or sandy soils

derived from ancient sandy coastal deposits. San Francisco lessingia is now

narrowly associated with either sparse vegetative cover or substantial vegetation

gaps, usually related to past artificial disturbance of the substrate or the

vegetation. Historic populations were probably associated with early stages of

succession following natural dune blowouts, or other local disturbances within

coastal dune scrub. The species may also have occupied stable dune vegetation

kept sparse or open by nutrient or drought stress. Spence (1964) observed that

annual Lessingia species were typical of dry, open, and somewhat disturbed

habitats, occurring in small, dense, disjunct populations. Nearly all historic

collections of San Francisco lessingia with specific locality and habitat

information have been made in areas with disturbed dune deposits. Exceptions

include one collection (Raven [1954], CAS 491875, Table 1) that was obtained

from weak, eroding sandy bluffs (Colma Formation), and the historic Ocean View

locality (Table 1; Dudley [1899]), which is mapped as Colma Formation sands as

well (Bonilla 1965). The Hillside Park (Daly City) population also occurs on

Colma Formation sand deposits that have been disturbed and are subject to

reworking by wind and runoff erosion. There are no historic or modern records of

San Francisco lessingia occurrence in foredunes or similar youthful dune areas

with little or no soil development and strong dune mobility. 

Undisturbed areas of dune scrub or nonnative grassland with high density and

cover of vegetation show little tendency to become invaded by San Francisco

lessingia, even when they are adjacent to source populations of this species (P.

Baye, pers. observ. 1997 to 1998). In contrast, San Francisco lessingia can spread

vigorously under favorable climate conditions when extensive substrate

disturbance occurs. After dune restoration sites at Lobos Creek and Wherry

dunes were graded, the low density and cover of vegetation encouraged rapid

spread of San Francisco lessingia from 1997 to 1998 (J. Cannon, pers. comm.

1997, Golden Gate National Recreation Area unpublished data 1998). Within San
38

Francisco lessingia populations at Lobos Dunes and behind the former Marine

Hospital [Public Health Services Building], plant size and fecundity appear to

increase markedly at the edges of vegetation gaps, such as footpath margins,

where potential competition is reduced (P. Baye, pers. observ. 1998). The species

is intolerant, however, of even a few centimeters (an inch or less) of active sand

accretion during its seedling, juvenile, and mature developmental phases (Pogge

1998). 

Based on modern and historic evidence, it appears likely that prior to European

settlement San Francisco lessingia was a sporadic local colonist of secondary

blowouts during their restabilization phase, or around their vegetated margins.

San Francisco lessingia probably persisted at sites where dune scrub and

grassland vegetation remained sparse or open, especially with patches of exposed

but relatively immobile bare sand. Analogous habitat conditions can be observed

today in relatively small secondary blowouts within well-stabilized portions of

large dunes on the central California coast. In the stabilizing phases of secondary

blowout succession, when bare sand begins to stabilize under the influence of

recolonizing plants, vegetative cover is sparse and low, but prevailing rates of

sand accretion or deflation are also reduced to low levels (P. Baye, pers. observ.

1990 to 1999). San Francisco lessingia would likely have been an element of this

vegetation, and probably persisted around wind-scarped edges of blowout crests

where chronic, low-level erosion gaps were sustained. Under modern conditions,

disturbances such as trampling, earthmoving, and grading probably serve as

surrogates for dune blowouts.

b. Plant Associations. San Francisco lessingia is associated with a diverse range

of species typical of dune scrub and grassland in San Francisco, as well as many

nonnative species (Appendix I). Spence (1964) observed that Lessingia species

are generally unable to grow and compete where the associated vegetation is

dense. Accordingly, some associations between San Francisco lessingia and other

species (particularly shrubs) are likely to be antagonistic at small spatial scales. 

However, positive correlations may be expected among herbaceous species with

similar life-history, low growth habit, or colonizing traits, which occupy

vegetation gaps and blowouts between patches of dense shrubby vegetation. 

Low-growing herbaceous species associated with San Francisco lessingia today

include dune gilia (Gilia capitata ssp. chamissonis), San Francisco spineflower
39

(Chorizanthe cuspidata), dune evening-primrose (Camissonia cheiranthifolia),

annual dune evening-primroses (Camissonia micrantha and Camissonia

contorta), annual plantain (Plantago erecta), coast fiddleneck (Amsinckia

spectabilis), sand mat (Cardionema ramossisima), dune popcorn-flower

(Cryptantha leiocarpa), and annual phacelia or wild heliotrope (Phacelia

distans). 

Dune shrub species are probably positively associated with San Francisco

lessingia at a coarse scale (large sample areas with shared overall habitat), but

negatively associated with it at a small scale (different microenvironments within

habitats). Relatively tall or coarse perennial herbaceous plant species associated

today with San Francisco lessingia (possibly in later stages of succession) include

dune goldenrod (Solidago spathulata; at Daly City, in a small relict population

above Baker Beach, and at two restoration sites at the Presidio), California

phacelia (Phacelia californica; Daly City only today), yarrow (Achillea

millefolium), bracken fern (Pteridium aquilinum), and California poppy

(Eschscholzia californica). Growth, survivorship, and reproduction of San

Francisco lessingia at the Presidio are inhibited by competition from nonnative

grasses, particularly vulpia (Vulpia spp.) and ripgut brome (Bromus diandrus)

(Pogge 1998, Brian O’Neill, National Park Service, in litt. 2002). The National

Park Service also notes that vulpia is relatively harder to remove by hand than

ripgut brome. San Francisco lessingia also occurs with other invasive nonnative

annual weeds such as foxtail (Vulpia bromoides), horseweed (Conyza

canadensis), and wild oat (Avena fatua) (P. Baye, pers. observ. 1997 to 1998). 

c. Life History and Reproduction. Seedlings of San Francisco lessingia emerge

from late fall to spring, soon after periods of rainfall and increased near-surface

sand moisture (Pogge 1998, J. Cannon, pers. comm. 1996). Seedlings emerging

at different times and growing at different rates form a size hierarchy. Individuals

that emerge and achieve large size earlier in the growing season have higher

reproductive output (Pogge 1998). There are two marked growth phases in

development (Howell 1929). During the rainy season seedlings develop into

juveniles, which are unbranched vegetative short plants (Figure 4), typically with

basal rosettes (relatively broad leaves on short erect stems with the growing tip

near the ground). Around the end of the rainy season (mid- to late spring) the

erect central shoot of the juvenile plants elongates, then branches profusely,
40

producing smaller leaves. The plant develops a low, spreading, bushy growth

habit in open conditions before it enters reproductive phase (Figure 4, Howell

1929, Pogge 1998). Branching continues repeatedly after flowering and seed set. 

Lateral shoots develop below individual flowerheads and seedheads, terminating

in new clusters of flowerheads. These flowerheads in turn develop more lateral

shoots below them, enabling flowering and seed production to occur continuously

together to almost to the end of the life-cycle (P. Baye, pers. observ. 1998). 

Flowerheads begin to form as early as May; unopened flowerheads are common

in early summer. Flowers usually open beginning in mid- to late summer, with

abundant flowering in August-September, continuing through November (Howell

1929, Pogge 1998). Reproductive output of individual plants is highly variable. 

Pogge (1998) found a range from 1 to over 1,400 flowerheads per plant,

depending on plant size. Large plants in sparsely vegetated areas may produce

many hundreds of flowerheads, each bearing up to 40 florets (potential seeds;

Lane 1993), but actual average number of seeds per flowerhead is about 26

(Pogge 1998), implying that the largest individuals may produce up to 36,400

seeds. Individuals competing with dense nonnative annual grasses or growing in

drier microsites may be very short and sparsely branched, with few or tens of

flowerheads. 

Spence (1964) performed artificial hybridization experiments with many 

Lessingia species, and determined that Lessingia species are generally selfincompatible (infertile when restricted to self-fertilization). Sibling plants grown

in isolation from parent populations sometimes fail to produce viable seed despite

availability of potential pollinators (P. Baye, pers. observ. 1998), which suggests

self-incompatibility may limit natural establishment of isolated small founder

populations established from infrequent long-distance dispersal events. Artificial

interspecific hybrids develop rather easily in cultivated populations (Spence

1964). Natural interspecific hybrids of San Francisco lessingia, however, would

not be expected to occur because the species is geographically isolated from other

species in the genus (Spence 1964).

Pollination may be achieved by insects, or possibly by wind as well (Spence

1964). Pollen of Lessingia species is light and dry (Spence 1964) rather than

heavy and sticky, as would be expected for flowers exclusively adapted to insect
41

pollination (Faegri and van der Pijl 1979, Proctor et al. 1996). Some windpollinated grassland species are also pollinated by insects (Faegri and van der Pijl

1979). Spence (1964) failed to detect any indication of pollen transfer by insects

in any wild populations of Lessingia species, even though potential pollinators

(syrphid flies and other Diptera, which lacked detectable pollen traces on their

bodies) were present. Other potential insect pollinators observed visiting

Lessingia germanorum flowers include numerous moths and butterflies (order

Lepidoptera), such as snout-nosed moths (family Pyralidae), skippers

(Hesperidae), ringlets (Satyridae), cabbage whites (Pieridae, Pieris rapae), blues

and hairstreaks (Lycaenidae), and American Ladies, (Nymphalidae, Vanessa

virginiensis); flies (order Diptera); bees and wasps (Hymenoptera), including

sand wasps (family Sphecidae), sweat bees (Halictidae), andrenid bees

(Andrenidae), bumblebees (Apidae, Bombus vosnesenskii); plant bugs (order

Hemiptera, family Miridae); and weevils (order Coleoptera, family

Curculionidae) (A. Whelchel unpublished data 1998). 

Ripe achenes (plumed “seeds”) begin to disperse in September and continue

through late fall. Earliest achenes have been observed in late June (Pogge 1998). 

Achenes are primarily wind-dispersed, as indicated by their light weight, small

size, and well-developed pappus (Spence 1964). Seed set of populations at the

Presidio appears to be consistently high (P. Baye, pers. observ. 1996 to 1998). 

Seed dispersal distance has not been studied, but seedling distribution tends to be

contagious around parent plants. Landscape barriers to dispersal, such as tree

plantations, may be more significant barriers to dispersal than inherent dispersal

ability of achenes. Seeds may also possibly be passively dispersed by humans, by

adherence of seed to footwear or clothing (J. Cannon, pers. comm. 1997). While

dispersal ability of San Francisco lessingia may be low because of landscape and

habitat constraints, its colonizing ability in suitable open or disturbed sandy

vegetation gaps (e.g., Hillside Park, Lobos Dunes, Wherry Dunes) appears to be

quite strong.

San Francisco lessingia is easily cultivated and shows no special requirement for

soil composition (Spence 1964). Germination also occurs without difficulty in

cultivation. Seeds lack innate, physiological dormancy (Spence 1964). Seeds

germinate very near the sand surface; as little as 1 to 2 millimeters (less than 0.08

inch) of sand burial strongly inhibits germination (Pogge 1998). Stored seeds
42

retain viability for at least 5 years with only a 50 percent reduction in the ability

to germinate; 2-year-old seed can exhibit 95 to 100 percent germination (Spence

1964). These germination data suggest that at least a short-lived soil seed bank is

likely to occur in nature, but the long-term viability of soil seed banks for this

species is unknown. The long-term viability of artificially stored refrigerated

seed is also unknown. Mollette (1998) concluded that San Francisco lessingia

formed only a transient seed bank, based on relatively short-term (2 years) field

data. Germination in cultivation occurs within 3 or 4 days after sowing and

watering, with a few seeds germinating as long as 127 days later (Pogge 1998). 

Seed sown in newly restored dune habitat in years of high rainfall resulted in high

seedling establishment (J. Cannon, pers. comm. 1997). Germination rate is

unaffected by the presence of ripgut brome under experimentally manipulated

field conditions. Germination percentage in cultivation may reach nearly 90

percent, and germination percentage in the field has been observed from over 43

to nearly 47 percent (Pogge 1998). 

4. Causes of Decline and Threats to Survival

San Francisco lessingia was listed as threatened (U.S. Fish and Wildlife Service

1997) (62 FR 33368) based on an analysis of the five factors under section 4(a)(1)

of the Endangered Species Act: (1) the present or threatened destruction,

modification, or curtailment of its habitat or range (urbanization, invasion of alien

plants, sand quarrying, bulldozing, and damage by pedestrians, bicycles, and offroad vehicles); (2) overutilization for commercial, recreational, scientific, or

education purposes (possible future threat); (3) disease or predation (no known

threat); (4) the inadequacy of existing regulatory mechanisms (limited protection

from State law and National Park Service land management); and (5) other

natural or manmade factors affecting its continued existence (bicycles,

pedestrians, off-road-vehicles, garbage dumping, fertilizer-contaminated runoff,

and habitat fragmentation). Habitat loss and adverse alteration of ecological

processes are the principal historic causes of decline of San Francisco lessingia. 

The resulting small numbers and localized distribution of the species have in

themselves become a primary threat to survival because of the amplified risk of

extinction through random catastrophic events or progressive vegetation change. 

Other threats identified above are relatively more localized or secondary,

dependent aspects of the dominant threat of habitat loss and degradation.
43

The initial historic cause of decline in the abundance and distribution of San

Francisco lessingia was the conversion of patchy dune scrub habitat to conifer

plantations, military posts, residential development, golf courses, and other urban

land uses. These conversions began at a significant scale quite early, around the

mid-19th century, and reached their maximum extent around the 1950's and

1960's when the last large tracts of privately owned undeveloped lands in San

Francisco were urbanized (Kaufeldt 1954). Land use conversion reduced

population size and distribution of dune plants to small remnants. These small

remnants were isolated from natural processes that may have maintained stand

dynamics, particularly secondary dune blowout succession (spontaneous

initiation, enlargement, and later revegetation of bare, wind-eroded patches in

sand dunes). 

The essential ecological processes that have been lost in San Francisco dune

vegetation are unimpeded transport of sand from beaches and mobile dune

complexes, and natural disturbances at both large and small scales that controlled

vegetation gap dynamics. Other essential ecological and geomorphic properties

of the San Francisco dune system that have been lost or diminished significantly

include variation in dune topography, which controls windflow patterns and

intensity (Olson 1958); burial and rejuvenation of stabilized dunes and dune

slacks (wet depressions) by mobile dunes; wind erosion and sand transport in

dune blowouts; and sand slope adjustment in eroding dune scarps (Jungerius et al.

1981, Carter 1988, Gares and Nordstrom 1990, Fraser et al. 1998). These

properties are typical of dune systems with complex vegetation mosaics

associated with development of blowouts. Blowout dynamics and associated

vegetation patterns develop over large spatial scales and appropriate landscape

positions, but not in small, isolated dune remnants within a sheltering matrix of

buildings and planted stands of nonnative trees. Homes and tree plantings create

obstacles to sand transport and create wind-shadows, reducing wind velocities

and potential sand transport corridors in remnant dune vegetation. They also

eliminate potential sand fetch areas — alignments along which dominant winds

may sweep sand to deposit downwind. These influences tend to promote

stabilization of small dune remnants, and encourage relatively complete closure of

dune scrub vegetation. Without recurrent disturbances and ecological

regeneration of vegetation gaps, and the characteristic physiological stresses of
44

coastal dunes that maintain integrity of the vegetation character, the long-term

persistence of San Francisco lessingia colonies is unlikely.

The artificial stabilization of dunes in San Francisco, and an artificial reduced 

disturbance regime in the dune remnants, would probably have caused extinction

of San Francisco lessingia if local artificial disturbance had not partially

compensated for them. Occasional earthmoving, grading activities, creation of

oversteepened erosional slopes, vehicle tracks, trampling, and other disturbances

caused by humans probably acted as surrogates for natural disturbances, and

maintained sandy vegetation gaps in coastal dune scrub that enabled San

Francisco lessingia to survive in small and unstable remnant patches for many

decades. When populations of San Francisco lessingia were relatively large and

well-distributed, artificial disturbances caused by humans were probably

beneficial to the survival of the species in altered remnant dune habitats. 

Disturbances within excessively small habitat remnants, however, may be

detrimental to the persistence of San Francisco lessingia today. Disturbances that

would otherwise be beneficial may jeopardize remnant populations that are

already precariously small. Whether disturbances are beneficial or harmful to

remnant populations depends on the scale of disturbance relative to population

size and distribution. Artificial disturbances are probably beneficial to San

Francisco lessingia when they are intermittent and patchy within an extensive

population. If disturbances are relatively frequent, chronic, or widespread, they

are likely to become detrimental to ecological regeneration of small populations

or small habitat patches. 

The secondary cause of decline, and the most important short-term threat to

survival of San Francisco lessingia, is habitat change caused mainly by the spread

of invasive nonnative vegetation, but also by excessive density and cover of

native woody shrub species, especially in sheltered conditions (such as planted

groves of evergreen trees). The principal nonnative invasive plants in older,

stabilized San Francisco dunes are iceplant (Carpobrotus edulis and its hybrids;

Vila and D’Antonio 1998), and annual Mediterranean grasses (in particular vulpia

and ripgut brome). Other species that are locally aggressive and colonial invaders

of stable dunes and sandy urban soils include an annual iceplant (Conicosia

pugioniformis); Bermuda-sorrel (Oxalis pes-capreae); and a relatively recently

established South African grass (Ehrharta erecta) (Appendix III). Species like
45

iceplant, vulpia, and ripgut brome tend to produce continuous, dense stands that

lack vegetation gaps and accumulate surface plant litter and soil organic matter. 

These conditions apparently discourage ecological regeneration of San Francisco

lessingia. 

Introduced trees from plantings in former dunes (Monterey cypress, Cupressus

macrocarpa; Monterey pine, Pinus radiata; blue gum; Appendix III) also spread

spontaneously into dune scrub over time, and may gradually convert dune scrub

remnants to nonnative “forests.” This threat is borne out by the fact that Presidio

natural resource managers remove several hundred Monterey pine, blue gum, and

particularly Monterey cypress saplings from dune restoration sites each year

(Brian O’Neill, National Park Service, in litt. 2002). Stands of nonnative trees,

whether planted or naturalized, have significant indirect impacts as well; the

wind-shadows of tall nonnative tree groves also further diminish the possibility of

natural disturbance regimes (dune blowout and slope dynamics) from

reestablishing, even in dune restoration sites. They can also rapidly reverse

potential gains in species recovery by degrading the quality of restored dune

habitat for San Francisco lessingia. Trees induce fog-drip moisture inputs to soil,

and tree canopy shading reduces evapotranspiration rates of the ground layer

vegetation. Fog drip and shade from trees reduce moisture stress that would

naturally exclude many shrubs and herbs from dune vegetation, and can promote

dense herbaceous undergrowth or dense mats of leaf litter even beyond their driplines, apparently inhibiting growth and spread of San Francisco lessingia (P.

Baye, pers. observ. 1996 to 1999). 

Potentially beneficial disturbances of San Francisco lessingia habitat may

nonetheless harm some existing populations, especially small ones. Trampling,

off-road bicycle or other vehicle use, small-scale sand quarrying, and burrowing

by dogs had been identified as threats to the species’ survival in the 1980's and

early 1990's (U.S. Fish and Wildlife Service 1997). In the Presidio, these impacts

are proportionally smaller today because of expanded habitat area, increased

public education, restricted pedestrian access to sensitive vegetation, and

symbolic fencing (fencing that is easily defeated physically, but defines areas of

restricted access). If the Golden Gate National Recreation Area is unable to

maintain dune vegetation quality, or becomes unable to adequately regulate
46

trampling impacts, these disturbances could again become significantly

detrimental to some colonies of San Francisco lessingia.

Small population size, small habitat patch size, and fragmentation (isolation) of

habitat patches in the urban landscape are indirect effects of habitat loss and

degradation. These indirect effects are probably independent threats to the

survival of San Francisco lessingia. When there are few suitable habitat patches

and seed dispersal distances to the patches are relatively large, local extinction of

San Francisco lessingia becomes likely. Population failure may also occur when

infrequent opportunities for new colonization (formation of sparse or bare

patches) are not favorably synchronized with local extinction of small colonies. 

Turnover of local colonies in dynamic patches is possible when there are

numerous, well-distributed populations within efficient dispersal distance of each

other (White 1996). In theory, such turnover would result in dynamically stable

populations, involving the local extinction of some patches, roughly balanced by

spontaneous establishment of new colonies in recently-disturbed sites. 

Dynamically stable populations of San Francisco lessingia do not remain near an

“average” population size but instead may fluctuate strongly over short time

scales (about 3 to 7 years). For this reason, fixed population size targets have

limited applicability for San Francisco lessingia.

Amelioration of natural environmental stresses inherent in coastal dunes (e.g., soil

moisture deficiency, soil nutrient deficiency, high evapotranspiration rates, and

mechanical wind-stress) is also likely to be detrimental to long-term survival or

recovery of San Francisco lessingia. Reduction of environmental stress or

enrichment of plant resources (nutrients, moisture) tends to increase growth and

density of vegetation, particularly weedy vegetation. Although San Francisco

lessingia individuals may grow well in such favorable environments, the species

is unable to ecologically regenerate well in dense vegetation. Therefore, addition

of irrigation or fertilizer to dune soils for landscape improvements, or shading and

fog-drip caused by planting of nonnative trees, tend to be antagonistic to annual

dune herbs like San Francisco lessingia. Planting of tall, invasive non-native tree

species or allowing such tree seedlings to establish in open sandy areas would

have similar detrimental effects on soil suitability for San Francisco lessingia. In

the long-term, addition of any water, nutrients, or organic soil amendments in

restorable potential San Francisco lessingia habitat could result in effectively
47

irreversible soil conditions (or conditions costly or difficult to reverse) that are

unfavorable for San Francisco lessingia’s recovery. Landscape maintenance or

enhancement is a potential conflict in some Presidio settings where restorable

habitat exists, and may be a problem at other restorable sites, as well. A

distinction is to be made between short-term watering and long-term watering

with use of soil amendments. Short-term watering may be necessary to establish

perennial plants during dune scrub restoration and of landscape species in

adjacent buffer areas. However, long-term and regular watering that increases

weeds and creates strong competition would result in reduced regeneration of San

Francisco lessingia, and may result in local extirpations.

The recent transfer of Golden Gate National Recreation Area lands of the Presidio

(other than the immediate coastline) to the Presidio Trust, a wholly owned

government corporation (16 U.S.C. § 460 bb appendix), has unclear implications

for the recovery of San Francisco lessingia. The Presidio Trust is currently

developing and leasing lands within the Presidio for commercial use (K.

Feyerabend, pers. comm. 1998). Landscaping for new or renovated buildings

could include ornamental plants or new weeds that could invade San Francisco

lessingia populations; trees that act as windbreaks, fog-drip collectors, or shade

sources; or use of irrigation, mulches, or fertilizer. These landscaping features

could indirectly affect San Francisco lessingia populations, habitat quality, or

habitat restoration potential. Development or infrastructure improvement of areas

in the Presidio with highly restorable former dune soils might impede or reduce

the potential for full recovery of San Francisco lessingia, particularly where the

lessingia is near sites with high potential for re-use (e.g., Public Health Services

Hospital, Wherry Housing) or improvements and expansion (e.g., golf course).

The Presidio Trust guidelines and the Presidio Vegetation Management Plan

indicate that no new (or targeted invasive ) weeds will be introduced as a part of

any landscaping efforts (Craig Middleton, in litt., 2002). While we trust that the

Presidio management will be careful to avoid new introductions of weeds and will

assiduously control any that get in despite their efforts, there remains some

possibility that new weed species could become established and compete with San

Francisco lessingia in dune scrub habitats. This possibility underscores the need

to ameliorate other threats as much as possible by restoring larger areas of dune

scrub habitat ecosystems.
48

The legislation authorizing the Presidio Trust also mandates the transfer of

Presidio Trust lands to the General Services Administration for “disposal”

(allowing for the possible transfer to private ownership, rather than reversion to

the National Park Service) if financial success criteria of the Trust are not met

after a specified period of time. This impetus to meet financial criteria may

increase pressure to redevelop lands of the Presidio, and could limit the

availability of land needed for dune restoration for recovery of San Francisco

lessingia. If financial objectives of the Presidio Trust are not met, disposed

“privatized” lands could also significantly reduce opportunities for habitat

restoration and reintroduction of San Francisco lessingia. Disposal of Federal

lands to non-Federal jurisdiction would effectively remove much of the legal

protection for San Francisco lessingia and its habitat in the Presidio. These two

threats are not mutually exclusive, and could compound the loss of restorable

habitat otherwise available for recovery of the species.

Urban development of private lands (particularly the sandy steep slopes above

Bonnie Street, Daly City) could impede conservation of the Hillside Park

population of San Francisco lessingia. Residential development of the sand

slopes in the area would directly eliminate habitat and increase demand for

artificial stabilization (and thus habitat loss) on undeveloped portions of the

habitat. Urban runoff and subsurface seepage of fertilizer-contaminated irrigation

water may promote growth of dense vegetation (native or nonnative), which is

incompatible with persistence of dry, sparse, open San Francisco lessingia habitat. 

Further fragmentation and reduction in size of this small habitat patch would

diminish its potential long-term viability by reducing potential number and size of

subpopulations, reducing potential insect pollinator refugia, and increasing

urbanized edges.

Other potential threats, such as overcollection (amateur or scientific destructive

sampling), herbivory, pests, and diseases appear to have been negligible or minor

contributors to the decline and endangerment of San Francisco lessingia. Loss of

adaptive genetic variability is often a concern for some endangered species that

are artificially reduced to small, isolated populations, although there is only

limited evidence that colonization or persistence of plants are related to genetic

variability (Barrett and Kohn 1991). Demographic and environmental factors

may be more important than genetic factors in determining viability of most plant
49

populations (Guerrant 1992, Nunney and Campbell 1993). Even though San

Francisco lessingia has been reduced to very small populations in the recent past,

it appears to have remained adaptable enough to exploit new and altered habitat

vigorously, as indicated by very rapid population growth following habitat

restoration. The prevalence of high seed production in all remnant and expanded

populations (J. Cannon, pers. comm. 1997; P. Baye, pers. observ. 1998) suggests

that pollinator limitation or low fertility (such as mating compatibility problems)

have not recently been significant contributors to the species’ decline. Habitat

loss and degradation have been the main threats to the species.

5. Conservation Efforts

Brian O’Neill, General Superintendent of the Golden Gate National Recreation

Area, petitioned us under the Endangered Species Act to emergency-list San

Francisco lessingia as endangered on May 28, 1991. We proposed to list San

Francisco lessingia as endangered on October 4, 1994 (U.S. Fish and Wildlife

Service 1994b). The final listing of the species as endangered was published June

19, 1997 (U.S. Fish and Wildlife Service 1997). San Francisco lessingia was also

listed as endangered by the State of California in 1990 (California Department of

Fish and Game 1992). The recovery priority number for San Francisco lessingia

is 2C, indicating a species that has a high degree of threat and high recovery

potential and is in conflict with construction or other development projects (U.S.

Fish and Wildlife Service 1983a, 1983b).

Important conservation efforts for the survival of San Francisco lessingia during

the last 20 years include suppression of nonnative invasive weeds; protection

against destruction of populations by mowing, sand quarrying, and grading;

protection against development; habitat restoration and expansion; population

augmentation (seeding into restored habitat); annual monitoring; and land use

planning that preserves opportunities for habitat restoration and reintroduction. 

Conservation efforts by local citizens and conservation organizations prior to and

after listing of San Francisco lessingia were critically important to the species’

survival. They included labor-intensive recurrent manual weeding; lobbying

landowners and municipal governments in Daly City to avoid severe short-term

impacts to remnant populations; and lobbying for long-term land use planning in

the Presidio compatible with conservation of the species (U.S. Fish and Wildlife
50

Service file information 1989 to 1990; J. Cannon, P. Holloran and S. Smith, pers.

comm. 1997 to 1998). Daly City has been responsive to citizen requests to avoid

impacts from maintenance activities, and has cooperated with citizen-led manual

weeding activities (S. Smith, pers. comm. 1998). Daly City and private

landowners have also practiced benign minimal vegetation management of sandy

slopes that have supported San Francisco lessingia. More aggressive slope

stabilization plantings would potentially have eliminated the population.

The Golden Gate National Recreation Area, following the closure of the Presidio

as a military facility, has led the conservation of San Francisco lessingia. The

Golden Gate National Recreation Area has established the Habitat Restoration

Team, and subsequently, the Presidio Park Stewards, stewardship programs aimed

at coordinating volunteer labor to control nonnative vegetation at Presidio

population sites. In cooperation with The Presidio Trust and the nonprofit Golden

Gate National Parks Conservancy, it has significantly expanded habitat (dune

scrub) and population size of San Francisco lessingia at the Lobos Dune

restoration area from less than 0.2 hectare (0.5 acre) with a small colony to

approximately 5 hectares (13 acres) supporting an extensive, vigorous population. 

This restoration effort anticipated recovery actions in this plan, and will

contribute substantially to the recovery of the species, assuming appropriate

vegetation management is sustained. The Golden Gate National Recreation Area

also has monitored population sizes of San Francisco lessingia over time and

fenced off remnant populations on the Presidio to protect them from excessive

trampling. Joint, cooperative stewardship (weeding) programs run by the Golden

Gate National Recreation Area, the Presidio Trust, and the Golden Gate National

Parks Conservancy have improved habitat quality of San Francisco lessingia sites. 

The Golden Gate National Recreation Area also has facilitated and permitted

graduate student research on San Francisco lessingia populations, providing

important basic biological information on pollination ecology and plant

interactions. The Presidio Trust also has been participating in Area B restoration

and stewardship efforts (Craig Middleton in litt. 2002). (Following the creation

of the Presidio Trust in 1996, the Presidio was divided into Areas A and B. Area

A continues to be managed by the Golden Gate National Recreation Area of the

National Park Service; Area B is managed by the Presidio Trust. The San

Francisco lessingia Presidio Recovery Unit includes parts of both Areas A and B.)
51

The San Bruno Mountain Habitat Conservation Plan, adopted in 1983 under

section 10(a)(1)(B) of the Endangered Species Act (U.S. Fish and Wildlife

Service permit PRT 2-9818, expiring March 31, 2013) before San Francisco

lessingia was federally listed as endangered in 1994, does not include or

otherwise protect the Hillside Park, Daly City site that supports San Francisco

lessingia.

6. Species Recovery Strategy for San Francisco Lessingia

The recovery strategy for San Francisco lessingia is based primarily on protecting

and expanding the existing populations within native coastal dune scrub

vegetation, followed by active reintroduction and expansion of San Francisco

lessingia in unoccupied, restored or enhanced habitat within its historic range. 

Neither protection of existing populations nor restoration and reintroduction

projects would alone be sufficient to recover the species in the long-term. Merely

maintaining the existing relict populations, which are surrounded by degraded

vegetation that has been invaded by nonnative species, would risk failure in the

long-term. Management of small remnant populations within extremely reduced

habitat fragments would be equivalent to mere cultivation of the species. 

Maintenance of small relict populations in the foreseeable future, however, will

provide interim insurance against extinction. These populations can be used as

sources of stock for reintroduction in dune restoration projects. 

The best chance for recovery of a species with highly reduced natural habitat

remaining is through jointly protecting existing populations and establishing

restoration/reintroduction programs for the species recovery (Falk 1992). 

Restoration and reintroduction depend heavily on unpredictable chance events in

changing environments — uncertainties that are inherent in restoration of dune

vegetation and establishment of founder populations (Kutner and Morse 1996,

White 1996). Restoration and reintroduction projects for San Francisco lessingia

may have less predictable success than intensive maintenance of relict individual

populations in static conditions, but they also have much higher potential longterm “yield” for dynamically stable populations that may be less dependent on

chronic manipulation. Successfully restored San Francisco lessingia populations

and dune communities also are more likely to exhibit natural ecological diversity

that can shape ongoing evolutionary processes affecting the species.
52

Recovery of San Francisco lessingia requires that appropriate vegetation

composition, structure, and dynamics be established and maintained on suitable

dune topography. In particular, areas of diverse dune topographic relief and

exposure are needed for a mosaic of dune scrub and patches of bare or sparsely

vegetated dune sand (blowouts) in various stages of recolonization by native dune

scrub vegetation. These ecological requirements are analogous to those proposed

for managing biological diversity of some noncoastal dune systems (Lesica and

Cooper 1999). The ecological and physical processes that maintain these features

operate only at relatively large spatial scales compared with the small remnant

patches of San Francisco lessingia at the time of listing. Thus, these processes

require at least some large sites (many tens of hectares in suitable configuration

and topography) with suitable exposure, aspect, sand supply, slope, and

disturbance agents. Nearly all of the reserves proposed for the species’ long-term

recovery (see below) meet this need.

The conceptual site-specific designs of San Francisco lessingia reserves within

three recovery units (see Chapter IV, Comprehensive Strategy of Recovery

Actions, San Francisco lessingia recovery), were based on localities of historic or

relict populations, suitable restorable substrate, landscape position, and feasibility

of restoration and protection (landscape constraints). Reserve design was also

influenced by basic ecological understanding of dune systems in the central

California coast region, and limited historical information on San Francisco’s

former dunes. Recovery objectives for San Francisco lessingia focus on three

main geographic recovery units. The main recovery units (Figure 7) are: (1) the

Presidio Recovery Unit, including the northern cluster of populations in restored

dune complexes within the Presidio (Lobos-upper Baker Beach-Wherry-Public

Health Services Hospital area); and (2) the Southern Recovery Unit, including a

southwestern pair of reserves at Hillside Park (Daly City) and a large population

at the restored Fort Funston dunes. Hillside Park and Fort Funston together

comprise the Southern Recovery Unit. A third subsidiary Recovery Unit,

comprising smaller satellite populations rather than larger core populations and

reserves, includes reintroduced populations at remnant dunes of Sunset Heights

and Sutro Heights near the Cliff House (Figure 1). These sites are within the

historic geographic and ecological range of the species, but are not among the few

specific historic San Francisco collection localities for which site-specific names

were attributed prior to urbanization of western San Francisco. They are, 

54

however, among the only sites available with potential restorable and manageable

habitat for San Francisco lessingia. These satellite reserves help compensate for

the irretrievable loss of historic populations and variable dune environments in

heavily urbanized parts of San Francisco where no open space remains.

Populations and vegetation within the main San Francisco lessingia recovery units

must be restored in settings and locations that will support natural disturbance

dynamics that can maintain their long-term integrity without excessive

intervention. These sites should also be managed to support associated native

dune plant vegetation, including live oak, arroyo willow and plant species of

concern. Potential founder population sources of San Francisco lessingia are

limited to a cluster of sites in the Presidio (northwestern range of the species on

dune sand) and one remote site in Daly City (southern range of the species on

sand derived from Pleistocene sand deposits). Seed sources for reintroduction

should be selected from the most geographically appropriate remnant populations. 

Selection of seed sources should consider proximity to modern and historic

populations, potential dispersal patterns and edaphic attributes of reintroduction

sites (soil texture, mineral, and chemical composition in relation to plant growth). 

General principles of seed source selection for plant reintroduction are provided

by Guerrant (1992, 1996), Falk et al. (1996), and Lesica and Allendorf (1999). 

San Francisco lessingia is an outcrossing annual that was historically widespread

in the Presidio dunes. This history suggests that it would be appropriate to

establish founder populations from pooled seed collected in multiple modern

“populations” in the Presidio. However, because of the probable similarity

between them, founders for new populations at Fort Funston (at the southern end

of historic range, nearest the historic collection localities at Lake Merced and

Ocean View, and with Colma Formation sand deposits most similar to Merced

formation sands) should be obtained only from Daly City. If relevant new data

indicate otherwise, recommendations for this seed source selection should be

reconsidered. For example, data suggesting high among-population genetic

variation within recovery units or substrate specific adaptation may indicate that a

different seed source selection strategy is more appropriate. A different strategy

may also be indicated if mixed seed source populations have low reproductive

output or low viability, as could be caused by inbreeding or outbreeding

depression. Restoration and management actions should include reintroduction of

extirpated plant species of concern, and at least one potentially-associated
55

endangered species (Layia carnosa) at appropriate locations. Additional recovery

measures include monitoring, applied research, seed storage, and public education

and outreach.

B. Raven’s Manzanita (Arctostaphylos hookeri G. Don. ssp. ravenii P. Wells)

1. Description and Taxonomy

Raven’s manzanita is a taxonomically ambiguous shrub within the complex and

variable genus Arctostaphylos (manzanitas, bearberry). It was most recently

classified as Arctostaphylos hookeri ssp. ravenii (Wells 1968, 1993). Raven’s

manzanita is a prostrate to ascending evergreen shrub in the heath family

(Ericaceae; Figure 8). It was reported to grow less than 60 centimeters (2 feet)

tall in historic inland localities (U.S. Fish and Wildlife Service 1984), but the

single wild plant today grows nearly prostrate on an exposed coastal site. 

Raven’s manzanita lacks burls (lignotubers), specialized flattened trunk-like

structures that are adapted to rapid vegetative regeneration following fires. Its

leathery, evergreen, round to round-elliptic leaves are 1 to 2 centimeters (0.3 to

0.7 inch) long, and are isofacial (have the same type of surface on both sides). 

Flowers are urn-shaped to round, with five-lobed white to pinkish corollas 4 to 5

millimeters (about 0.25 inch) long, with ovaries (floral precursors of fruits

containing undeveloped seeds) lacking pubescence (Wells 1993). Flowers appear

from mid-winter (in mild winters) to mid-spring (P. Baye unpublished data, U.S.

Fish and Wildlife Service 1984). Fruits are tan or brownish, round, and berry-like

with thick pulp, containing 2 to 10 stony seeds. The flower stalks are densely

covered with fine woolly hairs. Prostrate stems in prolonged contact with the

ground are reported to develop roots (U.S. Fish and Wildlife Service 1984). Key

distinguishing characteristics of this taxon, based on the single surviving

individual plant, are the combination of prostrate juvenile growth habit, round to

round-elliptic isofacial leaves, and small flowers and fruits (Wells 1993).

California manzanitas have a rich and a complex taxonomic and nomenclatural

history (Wells 1991), owing in part to the large number and diversity of species,

widespread hybridization, polyploidy (multiplication of chromosome sets), strong

geographic variation (especially local intermediates and variants), and local

ecological specialization (Markos et al. 1999, Raven and Axelrod 1978, Wells 
56

Figure 8. Raven’s manzanita (Arctostaphylos hookeri ssp. ravenii). (a)

inflorescence; (b) flowering shoots; (c) site of the remnant clone of

Raven’s manzanita near the World War II Memorial, Presidio. 

Adjacent vegetation: Monterey pines (Pinus radiata, background);

coastal grassland (foreground); and prostrate blue-blossom

(Ceanothus thyrsiflorus, foreground).

a b

c
57

1998). Taxonomic relationships and ranks of Arctostaphylos taxa have been

variously interpreted and revised throughout the history of California botany

(Jepson 1925; Abrams 1951; Munz 1959; Roof 1976, 1978, 1980; Wells 1991,

1998). The taxonomic and nomenclatural history of Raven’s manzanita reflects

this complexity. Early floras (regional inventories of wild plants) covering San

Francisco treated Raven’s manzanita variously (and with some confusion) as

Arctostaphylos pumila Nutt. (Brandegee 1892, Greene 1894), Arctostaphylos

pungens HBK (Bolander 1863, Behr 1888, Brandegee 1892), or Arctostaphylos

hookeri (Jepson 1925). More recent floras treated it as Franciscan manzanita,

Arctostaphylos franciscana (Munz 1959, Thomas 1961) or Arctostaphylos

hookeri ssp. ravenii (Wells 1993), which was disputed by Roof (1976, U.S. Fish

and Wildlife Service 1984). Roof (1976) considered it a distinct variety of the

variable and wide-ranging Mexican manzanita, Arctostaphylos pungens

(Arctostaphylos pungens HBK var. ravenii [Wells] Roof). 

Many botanists early in the 20th century treated Raven’s and Franciscan

manzanitas together as a single variable species. Raven, who rediscovered the

Presidio plant that later became the type of his namesake manzanita (Raven 1952,

Howell et al. 1958) originally treated Raven’s manzanita as an ambiguous variant

he referred with caution to Franciscan manzanita (Arctostaphylos franciscana

Eastw.). Franciscan manzanita is a rare endemic San Franciscan manzanita that

was entirely extinct in the wild at the time of Raven’s discovery, and now exists

only in cultivation (see Chapter III, Species of Concern). It was reclassified as

Arctostaphylos hookeri ssp. franciscana in Wells’ (1993) treatment of the genus. 

Raven (1952) also noted that his rediscovered Presidio manzanita was a close

match for an Eastwood specimen of Tamalpais manzanita (Arctostaphylos

montana Eastw. [= Arctostaphylos hookeri G. Don. ssp. montana (Eastw.) Wells])

from the type locality, Mt. Tamalpais (CAS 38507). Alice Eastwood, an eminent

manzanita taxonomist (Wells 1991), also failed to distinguish Raven’s manzanita

as distinct from Franciscan manzanita, despite familiarity with multiple wild

populations (Roof 1980). The past interpretation of Raven’s manzanita as a

variant of heterogeneous Franciscan manzanita is also reflected in multiple

herbarium sheets identified as Arctostaphylos franciscana that consist of mixed

samples of Franciscan and Raven’s manzanitas (e.g., CAS 38903, UC 185733,

JEPS 9390, UC 185737, UC 581330; U.S. Fish and Wildlife Service 1984, Roof

1976; T. Daniels, P. Wells, herbarium label annotation in CAS specimens)
58

indicating that early botanists treated them as one entity. These two taxa have

different numbers of chromosomal sets (ploidy; n = number of chromosomes in

one set = 26 in Raven’s manzanita, indicative of doubled chromosome sets, or

tetraploidy, n = 13 in Franciscan manzanita, indicative of a single chromosome

set, or diploidy; Wells 1968). 

In a recent taxonomic treatment of California manzanitas (Wells 1993), Raven’s

manzanita is distinguished by the character combinations of round leaves,

prostrate growth habit that persists in cultivation, small fruits 4 to 5 millimeters

(0.15 to 0.19 inch) wide, and nearly spherical flowers 4 to 5 millimeters (0.15 to

0.19 inch) long. By comparison, the sympatric (occurring in the same geographic

areas) Franciscan manzanita has oblanceolate leaves (longer than wide, wider

toward the tip), larger and reddish fruits 6 to 8 millimeters (0.24 to 0.32 inch)

wide, and larger urn-shaped corollas 5 to 7 millimeters (0.2 to 0.28 inch) long

(Roof 1980, Wells 1993). Other distinctions between these two taxa are

discussed by Roof (1976, 1980; U.S. Fish and Wildlife Service 1984). Tamalpais

manzanita is distinguished from Raven’s manzanita by its elliptic, rarely round

leaves (Howell 1949) (typically round-elliptic to round in Raven’s manzanita),

larger fruits (6 to 8 millimeters [0.24 to 0.32 inch] wide), and more woolly

inflorescence axis. Wells (1993) used the prostrate or matted habit of the one

remaining Raven’s manzanita plant as a general diagnostic trait of the entire

taxon. The importance of this trait was disputed by Roof (1976) because of

doubts about whether it was inherent species-wide trait, or a circumstantial trait of

the one remaining plant on a coastal headland environment. Behr (1892)

observed the former (1850's) San Francisco occurrence of the “tall form of

Manzanita still growing so abundantly on the slopes of Tamalpais” (presumably a

reference to the manzanita later named Tamalpais manzanita [Arctostaphylos

montana]), located at the site of the Protestant Orphan Asylum near Haight and

Laguna Streets. Given the similar appearance of some specimens of Tamalpais

manzanita and Raven’s manzanita (Raven 1952, Roof 1976), Behr likely

observed either former erect types of Raven’s or possibly Franciscan manzanita. 

No specimens of the Orphan Asylum manzanita were collected before the

vegetation there was destroyed. Roof (1976) also observed that former mature

populations of Raven’s manzanita had either ascending or erect growth habits in

some sheltered interior San Francisco locations. Roof (U.S. Fish and Wildlife

Service 1984) therefore questioned using the prostrate growth habit of the
59

Presidio clone as a diagnostic trait for the species as a whole. Prostrate and

ascending growth forms of some low-growing manzanita species, such as the

related Tamalpais manzanita, may be attributable in part to environmental factors

such as exposure (Howell 1949), maturation (loss of prostrate juvenile growth

habit), or to heritable variation in growth habit within the taxon. Historical

accounts and herbarium specimens overall suggest that the population of Raven’s

manzanita formerly exhibited variability in growth habit and vegetative

morphology that is now lost. Substantial variability in these traits within

California manzanita species is not uncommon (Wells 1993).

Roof (1976, U.S. Fish and Wildlife Service 1984) speculated that Raven’s

manzanita may have been a derivative of past hybridization between Tamalpais

manzanita and bearberry (Arctostaphylos uva-ursi) a wide-ranging northern

species that today still occurs within the Bay area at Point Reyes and San Bruno

Mountain (Howell 1949, Roof 1976, U.S. Fish and Wildlife Service 1984). 

Tamalpais manzanita occurs almost exclusively on serpentine substrates, but

occasionally grows on bare sandstone (Howell 1949, Roof 1976). One particular

specimen of Tamalpais manzanita collected from Mt. Tamalpais by Alice

Eastwood in 1903 (CAS 35807) was found by Raven (1952) and Roof (U.S. Fish

and Wildlife Service 1984) to be very similar to Raven’s manzanita, but this form

has not been located in recent decades for comparison under modern taxonomic

treatment. Other collections of Tamalpais manzanita also have rounded leaves

like Raven’s manzanita (L. Rowntree, CAS 187995; Abrams 5610, DS 68655; P.

Baye, pers. observ. 1999). Roof considered the round-leaved Eastwood specimen

of Tamalpais manzanita to be “scarcely distinguishable from the Masonic

Cemetery form” of Raven’s manzanita (extirpated, lacking voucher specimens)

that he personally observed. Behr (1892) also identified the San Francisco

manzanita at the former Orphan Asylum site to be a “tall form” of Tamalpais

manzanita; this statement is reasonably interpreted as a reference to the similarity

between Tamalpais manzanita and an extinct upright variant of Raven’s

manzanita.

As interpreted by Wells (Wells 1968, 1993), Hooker’s manzanita (Arctostaphylos

hookeri) consists of a series of five rare subspecies distributed in sandy or rocky

coastal environments from Monterey to Marin County. All but subspecies ravenii

and franciscana (= Arctostaphylos franciscana Eastw.), which historically
60

occurred together in mixed populations in San Francisco, are geographically

isolated from one another. Two taxa, ssp. hearstiorum and ssp. hookeri, are

clustered in coastal Monterey and San Luis Obispo Counties, and are found on

sandy soils. Three taxa, (subspecies or species) ravenii, montana, and

franciscana, are clustered around the Golden Gate, and are associated primarily

with serpentine bedrock outcrops. The extensive sympatry (co-occurrence in the

same geographic area) and lack of ecological differentiation of the two San

Francisco endemic manzanitas does not fully fit the concept of the subspecies

currently recognized by the International Code of Botanical Nomenclature, which

is based on the concept of DuRietz (1930). This subspecies concept is based on a

“more or less distinct regional facies of a species,” a distinct geographical or

ecological race. Nonetheless, the different ploidy levels indicate that these two

entities presumably could not freely interbreed as a single population of one

species, despite their shared geography, ecology, and structural similarity. 

Recent molecular genetic evidence (based on analysis of nucleotide sequences

from ribosomal DNA, genetic material from chromosomes of cell’s nucleus that

code for subcellular structures called ribosomes) suggests that Wells’ treatment of

Arctostaphylos hookeri, including Raven’s manzanita, does not comprise a

natural group (a unit of classification that reflects shared evolutionary descent)

(Markos et al. 1999). The genetic analyses of Markos et al. (1999) provides

evidence that the three manzanita taxa associated with serpentine soils of the Bay

area are related by common ancestry (a lineage including Californian A. uva-ursi

and A. tomentosa, or A. pungens, depending on the method of analysis). Their

analysis indicates that the two endemic San Francisco manzanitas are not closely

related to the southern subspecies of A. hookeri, which fall in a distinct lineage,

despite their overall similarity in some aspects of morphological appearance. 

Therefore, Wells’ subspecies of A. hookeri do not all share the same ancestors,

making the A. hookeri of Wells’ classification an “artificial” (paraphyletic)

group. The molecular genetic evidence also indicates that the pattern of evolution

in these taxa is complex, suggesting either past hybridization or sorting of

complex variable ancestors (Markos et al. 1999). 

The recent molecular genetic analysis of San Francisco manzanita relationships

(Markos et al. 1999) suggests that it is untenable to retain Raven’s manzanita as a

subspecies of Arctostaphylos hookeri in a natural classification, as Roof (1976,
61

1980) also earlier suggested on the basis of morphological and ecological

evidence. This analysis leaves the taxonomic status and nomenclature of Raven’s

manzanita once again unresolved and in need of revision. Markos et al. (1999)

concluded that it would be premature to revise the formal nomenclature and

taxonomy of San Franciscan manzanitas until more comprehensive genetic data

sets are available for other manzanita taxa. The conventional usage of Wells’

nomenclature, popularly used since its publication in the Jepson Manual

(Hickman 1993), the standard contemporary revised flora of California, is

followed in this recovery plan without prejudice to scientifically valid taxonomic

rank or placement of Raven’s manzanita.

Raven’s manzanita was originally distinguished as a taxonomically distinct entity

based on classical morphological methods and chromosome counts (Wells 1968). 

These methods alone cannot adequately distinguish among “pure” species,

hybrids, introgressants, or stabilized hybrid species (reproductive stable

populations derived from ancestral hybrid population, unlike sterile or transient

unstable hybrids) (Rieseberg 1991), including manzanitas (Ellstrand et al. 1987).

Indeed, the idea of a “pure” species in Arctostaphylos, with its many poorly

defined taxa and prevalent hybridization (Wells 1968, 1993; Roof 1976) has often

been difficult to apply over the history of taxonomic work in the genus (Wells

1991). Additional research, based on more intensive geographic sampling and

molecular genetic analysis, is needed to explore the ancestry of Raven’s

manzanita and its relationship with the other similar and related manzanitas

occurring near the Golden Gate: A. montana, A. franciscana, and wider-ranging

species like A. uva-ursi, A. tomentosa, and A. pungens. Whatever the taxonomic

rank and placement of Raven’s manzanita, its endangered status and conservation

priority remain the same: Holsinger and Gottlieb (1991) argue that stabilized

hybrid/introgressant populations are as worthy of conservation as “pure” named

taxa.

2. Current and Historic Distribution

Only a single natural clonal colony of Raven’s manzanita remains on an oceanfacing serpentine bedrock outcrop within a larger serpentine soil area near the

World War II Memorial at the end of Kobbe Avenue in the Presidio, above Baker

Beach, San Francisco. All other populations in San Francisco have been
62

extirpated. The clonal colony, rediscovered around 1950, is probably well over a

century old (Raven 1952). This population of one plant has been augmented by

planting artificially propagated genetically identical daughter clones of the mother

plant in its vicinity. Additional genetically identical populations have been

established by transplanting clonally propagated (rooted cuttings) plants in the

general vicinity of this site at three nearby locations. A clone also has been

planted at another serpentine outcrop at Inspiration Point in the Presidio, off

Arguello Boulevard. This same clone also has been maintained in cultivated

populations in San Francisco and Berkeley. Unlike Franciscan manzanita, no

plants of Raven’s manzanita were salvaged from former interior San Francisco

localities before they were destroyed by urban development. Raven’s manzanitas

at interior sites may not have attracted conservation attention because they were

once considered to be atypical forms of Franciscan manzanita (Roof 1976). The

clone planted at Inspiration Point has not been seen for about 5 years.

Historic San Francisco manzanita localities that supported both Franciscan

manzanita and Raven’s manzanita included: (1) the former Laurel Hill Cemetery;

(2) the former Masonic Cemetery (near Lone Mountain; the “base of Lone

Mountain” locality of “Arctostaphylos pumila Nutt.” reported by Greene (1894)

may have been the Masonic Cemetery locality or a nearby population); (3) Mount

Davidson, in the south-central part of the City; and (4) the Presidio locality still

surviving (Figure 2). In addition, there is a record of “Arctostaphylos pumila”

(Behr 1892; a misnomer for either Franciscan or Raven’s manzanita, or perhaps

both) at the former Protestant Orphan Asylum (Laguna at Haight Street, long

urbanized in the late 1800's). Nearly all historic localities of creeping manzanitas

in San Francisco were outcrops of serpentine (all sites except Mount Davidson,

which comprises greenstone and mixed Franciscan rocks), which suggests limited

historic and prehistoric distribution and only local abundance. Evidence for

historic mixed populations consists of inadvertent inclusions of Raven’s

manzanita material within herbarium collections of Franciscan manzanita (see

Description and Taxonomy on page 56) and direct observations of co-occurrence

(Roof 1976, U.S. Fish and Wildlife Service 1984). Otherwise, definitive historic

records and surveys of Raven’s manzanita are lacking because it was not

recognized as distinct from Franciscan manzanita until long after all but one of its

populations were extirpated. 
63

It is possible that the limited historic records under-sampled and underrepresented the early historic abundance and distribution of Raven’s manzanita. 

Many collectors of Franciscan manzanita may have consciously or unconsciously

selected material most similar to the type of this taxon (Roof 1976), which could

have biased the sampling of specimens. Behr (1892) described “Arctostaphylos

pumila” (not the species endemic to sandy soils near Monterey, but the “creeping

manzanita” of Brandegee’s San Francisco flora of 1894) as “once abundant” in

his botanical memoirs of San Francisco in the 1850's, and predicted its extirpation

there nearly a century ago. It is unknown how many populations of San

Franciscan manzanitas (mixed Raven’s and Franciscan manzanita) occurred

during and prior to the 1850's. Herbarium collections of manzanitas in San

Francisco were made in the 20th century, after urbanization was well advanced in

much of the City. 

One other manzanita species, Arctostaphylos tomentosa (Pursh) Lindley ssp. rosei

(Eastw.) P. Wells (= Arctostaphylos crustacea Eastw. var. rosei [Eastw.]

McMinn; shaggy-barked manzanita or rosy manzanita; colloquially,

“Brotherhood Way manzanita”), an erect shrub, occurs naturally in southern San

Francisco east of Lake Merced on sandy Colma Formation deposits. The nearest

populations of other manzanita taxa growing in the wild (including one bearberry,

A. uva-ursi) are on San Bruno Mountain, San Mateo County (McClintock et al.

1990). Bearberry may have occurred in San Francisco during the Pleistocene or

earlier in the Holocene epoch (M. Vasey, pers. comm. 1999).

3. Ecology and Reproduction

The available information on the ecology of Raven’s manzanita is limited. Direct

ecological data include: (1) the records of mixed populations of the endemic San

Francisco manzanitas; (2) the restriction of historic localities to bedrock outcrops,

primarily of serpentinite; and (3) associated plant species, soil conditions, and

microclimate observations at the existing natural locality. Indirect or comparative

ecological data include information on similar taxa within Arctostaphylos. 

Generally, available data suggest that Raven’s manzanita is a slow-growing,

stress-tolerant evergreen shrub that is able to grow on serpentine soils with sparse

competing vegetation, but like many manzanitas (Gottlieb 1968, Kruckeberg

1977), is relatively intolerant of competition (especially shading from shrub or
64

tree canopies). It appears to have been locally abundant (Behr 1892) in relatively

isolated, localized open serpentinite outcrop colonies, determined by structural

geology of the local landscape. Serpentine rocks in San Francisco are restricted

to two broad shear zones (bands of rocks deformed and crushed by lateral fault

movements): the City College shear zone, and the Hunters Point shear zone

(Schlocker 1974, Figure 2).

The degree to which Raven’s and Franciscan manzanitas depend on serpentine

soil chemistry is questionable. All recorded populations, except the Mount

Davidson collections on greenstone, occurred on serpentine substrates, which is

circumstantial evidence that they are at least facultative serpentine taxa. 

However, both Raven’s and Franciscan manzanita have been successfully

cultivated for many years under nonserpentine, irrigated garden soil conditions

without growth abnormalities or indications of nutrient deficiencies (Tilden Park,

Berkeley; U.C. Berkeley Botanical Gardens, and Strybing Arboretum; McCarten

1986; D. Mahoney and H. Forbes, pers. comm. 1999). The related and

ecologically similar Tamalpais manzanita occurs principally on serpentine, but

also thrives on bare sandstone (Roof 1976). The successful growth of Raven’s

manzanita and related taxa on nonserpentine substrates at multiple locations

indicates that they have no specific physiological (nutritional) requirement for

serpentine soil chemistry per se. This result is consistent with the hypothesis that

many instances of serpentine plant endemism (narrow habitat and geographic

restriction) are based on the intolerance of competition, and tolerance of harsh

serpentine soil conditions that inhibit competition (Kruckeberg 1984), combined

with maintenance of sparse, open vegetation. Most other species that occur

frequently on serpentine soils in San Francisco (Appendix II) also occur on thin,

rocky or clayey soils derived from other types of bedrock. 

Soil analysis of the Presidio Raven’s manzanita site (McCarten 1986) and other

serpentine outcrop sites of the Presidio, confirm that these soils are rich in

magnesium, relatively low in major plant nutrients, and are mildly acidic. Soil

nitrogen in shallow surface samples (0 to 5 centimeters [less than 2 inches] depth)

of Presidio soils, however, were consistently higher than inland Marin County

serpentine sites sampled. No data are available on other San Francisco serpentine

soils outside the Presidio.
65

The native plant species associated with Raven’s manzanita at the Presidio site

(Appendix II below; Appendix 3, U.S. Fish and Wildlife Service 1984) have not

changed substantially since the 1984 recovery plan was prepared. Recent

additional observations indicate that the wild clone is intermixed with grasses and

forbs. The most frequent species interspersed within the manzanita clone include

Torrey’s melic-grass (Melica torreyana), miner’s lettuce (Claytonia perfoliata),

and soap plant (Chlorogalum pomeridianum). Species marginally associated with

the manzanita clone within low-growing grassland vegetation include red fescue

(Festuca rubra), junegrass (Koeleria macrantha), sedges (Carex tumulicola,

Carex spp.), purple needlegrass (Nassella pulchra), California oatgrass

(Danthonia californica), and goldfields (Lasthenia californica) (P. Baye

unpublished data 1998). These herbaceous species may have increased in

abundance since shrub and tree removal to protect the plant was completed in the

1980's. San Francisco owl’s-clover (Triphysaria floribunda) and paintbrushes

(Castilleja affinis ssp. affinis and Castilleja subinclusa ssp. franciscana) have not

recently appeared at the site of the remnant Raven’s manzanita clone, but were

evident in the past (U.S. Fish and Wildlife Service 1984, Howell et al. 1958). 

The endangered Presidio clarkia (Clarkia franciscana) population has persisted

since its introduction (seeded into the site by J. Roof in 1972; Roof 1972),

primarily where steep bedrock outcrops at the surface and minimal soil

development is evident. A small population (50-100 individuals) of the Marin

dwarf flax (Hesperolinon congestum) occurs in the vicinity of the planted

daughter clones. The vegetation within and adjacent to the manzanita colony is

either low and herbaceous, or prostrate and woody (blue blossom, Ceanothus

thyrsiflorus). 

Climate and environment affect growth and form of Raven’s manzanita. High

rainfall appears to promote incidence of twig blight, but also appears to support

luxuriant growth later in the growing season (P. Baye unpublished data 1997 to

1998). Some plants from extirpated populations in sheltered conditions

apparently developed a more ascending to erect, but low, growth habit compared

with wind-flagged plants on exposed bluffs (U.S. Fish and Wildlife Service 1984;

also see discussion in Description and Taxonomy above). Prostrate and erect

growth forms of Tamalpais manzanita, controlled by sheltering and exposure, also

were noted by Howell (Howell 1949). Mature clones of the Presidio Raven’s

manzanita genetic individual growing at Tilden Botanical Gardens in warmer,
66

sheltered inland conditions remain strongly prostrate, indicating a strong genetic

component to the growth habit of the lone individual from the exposed coastal site

(P. Baye, pers. observ. 1998). Prostrate habit may also sometimes be partly a

juvenile trait. The marine Mediterranean climate of San Francisco is normally

frost-free, with rains largely restricted to fall, winter, and spring months most

years (U.S. Fish and Wildlife Service 1984). All historic localities of Raven’s

manzanita are influenced to varying degrees by persistent marine fogs that

depress summer temperatures and reduce intensity of sunlight exposure compared

with inland conditions, but there is strong local variation in climate in San

Francisco, influenced by topography. Cool fog and onshore winds are typically

persistent along the Golden Gate at the type locality of Raven’s manzanita, but

microclimates at historic localities vary from foggy but relatively wind-sheltered

(Masonic and Laurel Hill Cemetery) to intermittently foggy, warmer and

sheltered (Orphan Asylum), and intermittently foggy but wind-exposed (Mount

Davidson) (P. Baye, pers. observ. 1984 to 1999).

There are no scientific data on the breeding system of Raven’s manzanita, and

available evidence is unclear. Raven’s manzanita has been reported to be an

obligate outcrosser (M. Parker, pers. comm. cited in McCarten 1986), a crossfertilizing species unable to produce significant amounts of viable seed from selfpollination. Obligate outcrossing plants require at least two genetically

compatible individuals to reproduce sexually. The remaining isolated Raven’s

manzanita in the Presidio, however, has been observed to set seed spontaneously,

which suggests either some degree of self-pollination or very long-distance hybrid

cross-pollination. Cultivated and wild manzanitas alike readily hybridize among

species (McMinn 1939; Wells 1968, 1991; Roof 1976; Ellstrand et al. 1987). The

wild Raven’s manzanita plant, however, is reported to produce few mature fruits

and seed (U.S. Fish and Wildlife Service 1984). No mature fruits were observed

on the Presidio remnant clone, or its daughter clones, in 1998 or 1999, despite

abundant flowering and presence of bees during flowering (P. Baye, pers. observ.

1998 to 1999). In contrast, both self-pollinated and open-pollinated Raven’s

manzanita in cultivation at the University of California, Berkeley, have been

reported to produce abundant seed, with about 20 percent viability in both lots (40

percent of the fruits with at least one viable seed) in 1995 (H. Forbes, pers. comm.

1999). Tetraploid manzanita species (ones with double sets of chromosomes),
67

such as Raven’s manzanita (Wells 1968) generally have significantly lower

percentage seed set than diploid species (Kelly and Parker 1991).

There have been no reports of natural seedling establishment around the remnant

wild Raven’s manzanita or elsewhere since it was rediscovered in 1952. The

absence of seedlings may be due to a lack of viable seed, seed predation, or lack

of sufficient seedling microsites in the undisturbed vegetation around the single

natural plant, or possibly other factors. No data are available on the natural

germination ecology of Raven’s manzanita. Propagation of other California

species of manzanita often requires moist-chilling, scarification (mechanical

attrition) of seed coats, or treatments that mimic burns (Lenz 1956, U.S. Fish and

Wildlife Service 1984 and references within). It is possible that seed germination

of Raven’s manzanita would be stimulated by burns (Keeley 1987), as in other

manzanita taxa farther south on the San Francisco Peninsula (McClintock et al.

1990). Tamalpais manzanita has been observed to vegetatively regenerate from

seed following fire (herbarium sheet annotation, Eastwood 12980, CAS 128697). 

Generally, seed germination of manzanitas is slow and erratic (Lenz 1956), traits

consistent with persistent seed banks from which seedlings are recruited

following disturbances (Thompson 1992). Naturally occurring dormant seed

banks occur in other Californian manzanita species, both in fire-adapted species

(post-burn resprouting manzanitas with woody burls rich in vegetative

regeneration buds) or fire-sensitive species (regenerating only from seed after

burns; Kelly and Parker 1990). However, most studies indicate that most seeds

produced do not accumulate in soil seed banks, and many seeds are lost to

predation (Kelly and Parker 1990).

Nothing is known of the genetic structure of Raven’s manzanita prior to its

extreme decline to one individual. Roof (U.S. Fish and Wildlife Service 1984)

and some herbarium label authors observed layering (rooting of prostrate stems),

a form of clonal growth, but natural spread of the clone has so far not resulted in

multiple root systems (fragmented clones) of the old wild plant (P. Baye, pers.

observ. 1998).
68

4. Causes of Decline and Threats to Survival

Raven’s manzanita was listed as threatened (U.S. Fish and Wildlife Service 1979)

based on an analysis of the five factors under section 4(a)(1) of the Endangered

Species Act: (1) the present or threatened destruction, modification, or

curtailment of its habitat or range (past urban development, vulnerability to

management errors due to small population size, competition from other plants);

(2) overutilization for commercial, recreational, scientific, or education purposes

(possible collection); (3) disease or predation (none noted at time of listing;

fungal twig blight occurs in wet years and occasional irruptions of tussock moth

caterpillars [Family Lymantriidae] are known to defoliate plants); (4) the

inadequacy of existing regulatory mechanisms (limited protection under

California law; currently transfer of land to Presidio Trust or private ownership

may increase development pressure and reduce protections); and (5) other natural

or manmade factors affecting its continued existence (possible lack of pollinators

noted at time of listing; also lack of genetic diversity, low reproductive success,

and possible hybridization with other manzanitas).

The major cause of historic decline in Raven’s manzanita populations was the

irreversible elimination of its habitat by San Francisco’s urban growth. The lack

of seedling colonization in new habitat (possibly due to observed low

reproductive output or poor dispersal of seed to isolated patches of suitable

seedling habitat) appears to have prevented it from overcoming the adverse

effects of local habitat loss and fragmentation. Current threats to its survival are

partly due to inherent risks associated with the extreme reduction in population

size to a single clone (genetic individual), and partly due to external threats. The

external threats to the single Presidio individual that were described in the

original recovery plan continue to some extent today, with the exception of

shading by trees, which were removed about 1984. The small population of

replicate clones on the Presidio is also vulnerable to fire, landslides, accidental

injury by road maintenance or vegetation management activities, and vandalism

(U.S. Fish and Wildlife Service 1984). 

The principal contemporary threat to the persistence of the long-lived original

clone of Raven’s manzanita is competition (interception of light) by vegetation

that overtops the prostrate plant (U.S. Fish and Wildlife Service 1984). The most
69

significant potential competitors are trees that are not native to the San Francisco

Peninsula and can overtop and shade the remnant wild manzanita clone: Monterey

cypress, Monterey pine, and blue gum. These trees have been removed from the

immediate vicinity of the remnant clone (U.S. Fish and Wildlife Service 1984),

but reinvasion from abundant local seed sources remains a threat. Other invasive

nonnative plants in the vicinity of the manzanita site that represent potential

competitive threats include iceplants (Carpobrotus edulis and its hybrids),

myoporum (Myoporum laetum), plume acacia (Albizia lophantha), wattles

(Acacia spp.), jubata grass or “pampas grass” (Cortaderia jubata), Cape ivy

(Arctotheca calendula), cotoneaster (Cotoneaster sp.), and nonnative annual

grasses. Of these species, jubata grass is now among the most invasive species on

serpentine bluffs, scarps, and landslides below the manzanita preserve site. It has

proven to be highly invasive to disturbed sites, and produces abundant plumed

seed capable of long-distance wind-dispersal. Although jubata grass abundance

has recently been suppressed on the bluffs, it recolonizes readily and can grow

very rapidly even on serpentine soil (P. Baye, pers. observ. 1993 to 1999). One

native shrub species, a prostrate form of blue-blossom, also competes with

Raven’s manzanita to some extent (U.S. Fish and Wildlife Service 1984), but has

apparently co-existed with it for decades without causing progressive decline in

the clone. 

Ongoing rigorous vegetation management is needed to suppress reinvasion of

nonnative vegetation in the immediate vicinity of the preserved clone at the War

Memorial site and in the surrounding area, which is a source of nonnative plant

seed. Reinvasion will remain a potential threat to the site even if contemporary

invasion levels are low. Reducing the effort to suppress these invasive species,

even temporarily, would probably enable them to recolonize the preservation site,

and resume their threat to the manzanita clone.

In years of frequent and late rains, Raven’s manzanita develops relatively

extensive infections by a twig blight (called “black smut” in the original recovery

plan; U.S. Fish and Wildlife Service 1984) that causes leaf necrosis (tissue death)

and dieback of whole sectors of stems in winter and early spring (S. Farrell, pers.

comm. 1998). Up to 40 percent of individual clones may suffer dieback in a

mosaic pattern during winter months. Affected clones typically quickly recover

from blight-induced dieback during the subsequent growing season. Recovery
70

occurs mainly by overgrowth of dead sectors by vegetative shoots from adjacent

portions of the clone in spring and early summer (P. Baye unpublished data 1997-

1999). Twig blight is likely due to fungus-like disease agents such as Phomopsis

spp., which affect other species of manzanita as well (Lenz 1956). Cultivated

specimens of other native manzanitas at Strybing Arboretum, San Francisco, vary

in susceptibility to twig blight in rainy years. At Strybing, the Raven’s manzanita

clone is relatively susceptible to blight, even compared with Franciscan

manzanita. Cultivated clonal replicates of Raven’s manzanita in the drier, warmer

inland hills of Berkeley (Alameda County) exhibited no blight symptoms after

several wet years (P. Baye, pers. observ. 1997 to 1998). The disease organism

responsible for “Sudden Oak Death,” a new species of Phytopthora (a genus of

“water-mold,” parasitic algae that include root-rot, stem-rot, and potato-blight

diseases) also has been detected recently on at least one native member of the

heath family (madrone, Arbutus menziesii), and its host range is expanding

(Fimrite 2001). This virulent new plant disease has uncertain potential impact on

native manzanita species.

The long-term threat of disease to the wild Raven’s manzanita clone is uncertain,

but the old age of the wild clone suggests that it diseases have been in the past a

short-term, cyclic impact, not a progressively degenerative problem. Fungal

infection appears to be a greater potential threat to the survival of the smaller

daughter clones, which have proportionally more necrotic area when infected. 

The former variability in pathogen resistance within the entire original population

of Raven’s manzanita is unknown. Natural genetic variability in pathogen

resistance within populations is widespread among plant species (Burdon 1987),

and some variation may have been lost with former extirpated populations of

Raven’s manzanita. Lack of adequate levels of genetic variability in pathogen

resistance traits may make plant populations more susceptible to disease

outbreaks and increased mortality (Burdon 1987, Huenneke 1991). If more

virulent strains of twig blight or other pathogens infest the population, dieback

could imperil the single wild manzanita clone. Cumulative effects of infection or

cumulative increases in inoculum potential (accumulation of diseased leaf litter, a

source of spores for reinfection) could also threaten the single clone. Fungicidal

treatment of infected plants may be impractical because manzanitas also form

obligate associations with beneficial mycorrhizal fungi.
71

An irruption of tussock moth caterpillars (family Lymantriidae) discovered in

1999 affected most of the daughter clones of Raven’s manzanita. Defoliation

varied from about 5 percent to as much as 60 percent, and is believed to have

killed some of the daughters. The staff at the Golden Gate National Recreation

Area removed caterpillars by hand to reduce the herbivory. Infestation was much

reduced in 2000, and appears to have been absent since, but could recur. (Brian

O’Neill, National Park Service, in litt. 2002).

Raven’s manzanita’s evolutionary potential and its chances for survival in the

wild have probably suffered significantly because of its population crash to a

single genetic individual (the most extreme genetic “bottleneck” possible). This

situation poses a rare challenge for endangered plant species recovery (McMahon

1989, Falk 1992, Knapp and Connors 1999). Modern sexual reproduction of the

species appears to be lacking. Since the plant was rediscovered in 1952, no

seedling establishment has been detected, although the wild plant does set fruits

some years (U.S. Fish and Wildlife Service 1984) with some seed viability (H.

Forbes, pers. comm. 1998). The cause of the lack of seedling recruitment is not

known, but the challenges of reproduction by seed may be magnified by: (1) the

isolation of the plant from open, uncolonized habitat in a heterogeneous, patchy

serpentine environment; and (2) deficient adaptive morphological and

ecophysiological variation in seedlings produced by a single parent plant. A lack

of episodic environmental cues for germination (possibly fire or landslides)

around the site of the surviving wild plant may also constrain seedling

recruitment.

Recovery of endangered plants faces special problems in small populations

(Barrett and Kohn 1991), or populations represented by one (Knapp and Connors

1999) or two (Robichaux et al. 1997) genetic individuals. These problems

include failure by self-incompatible plants to produce viable seed and inbreeding

depression (decreased viability or fecundity [fitness] associated with mating

among relatives) (Barrett and Kohn 1991). Populations that typically outcross are

expected to be more vulnerable to inbreeding depression than populations with a

history of selfing (Barrett and Kohn 1991, Huenneke 1991). Strong inbreeding

depression, however, has been observed in some selfing species as well (Barrett

and Kohn 1991). Recovery could also be compromised by low levels of genetic

variation, especially in traits with adaptive ecological significance. Populations
72

with severely limited genetic variation are most vulnerable to extinction because

of reduced potential for evolution in response to environmental changes

(Beardsmore 1983, Huenneke 1991, Knapp and Connors 1999). Low levels of

genetic variation may also limit evolutionary changes necessary to cope with

reintroduction into environmentally variable portions of species’ historic ranges

(Frankel and Soulé 1981, Huenneke 1991, Lesica and Allendorf 1999), or

changes in pathogens or herbivores (Burdon 1987, Huenneke 1991). For

exceptionally rare plants like Raven’s manzanita, sexual reproduction in

populations with adequate levels of genetic variation may be important for

continued survival and evolution. Sexual reproduction among genetically diverse

individuals recombines (rearranges) genes, possibly creating new combinations of

traits that will be subject to natural selection (Crow 1988, Shields 1988).

Assimilation of a rare plant species by hybridization and introgression (repeated

backcrossing of a hybrid to one or both parental populations) with nonnative

species can also be a threat to survival (Rieseberg 1991, Ellstrand 1992, Levin et

al. 1996). “Pollen swamping” could occur if nonnative ornamental manzanitas

are planted in abundance near the War Memorial manzanita site in the Presidio. 

The distance from which pollinators are likely to transport manzanita pollen in

San Francisco is not known. Production of hybrid seed (with pollen parents of

nonnative manzanitas) could permanently corrupt any dormant soil seed banks

beneath the Raven’s manzanita clone, making natural seedling regeneration

unmanageable. This difficulty would also apply to any Raven’s manzanita

reintroduction sites.

Golden Gate National Recreation Area lands of the Presidio (away from the

immediate coastline) were recently transferred to the Presidio Trust, a wholly

owned government corporation (16 U.S.C. § 460 bb appendix). This change in

management has unclear implications for the recovery of Raven’s manzanita. 

One of the Presidio’s mandates is development and leasing of lands in the

Presidio for commercial use. In contrast, Golden Gate National Recreation Area

had no mandate or authority for commercial development of habitat or park lands. 

Development of potentially restorable habitat in derelict areas of the Presidio with

underlying serpentine rocks could reduce opportunities for restoration,

reintroduction, and recovery of Raven’s manzanita. Similarly, infrastructure

improvements associated with redevelopment (e.g., road widening, slope
73

stabilization, landscaping, recreational amenities, other facilities) could pre-empt

habitat restoration in undeveloped serpentine areas. The Trust does not merely

need to avoid impacts to existing Raven’s manzanita plants; it should provide

adequate priority and planning for future endangered species recovery, starting

with obtaining complete inventories of outcropping and near-surface serpentine

subsoils of the Presidio. 

An additional problem for endangered species conservation in the Presidio is that

the legislation authorizing the Presidio Trust mandates the transfer of Presidio

Trust lands to the Federal government’s General Services Administration for

disposal (allowing for the possible transfer of land to private ownership) if the

Trust fails to meet its financial success criteria after a specified period of time. 

Disposal of Federal lands to non-Federal jurisdiction would effectively remove

much of the legal protection for Raven’s manzanita and its habitat (actual or

potential) in the Presidio.

Recovery of Raven’s manzanita will depend on successful cooperation with

owners and managers of potential restoration and reintroduction sites, and their

neighbors. Cooperativeness may depend on public and institutional perceptions

of the burdens and risks of dealing with federally listed plant species. For

example, the environmental assessment for the Golden Gate National Recreation

Area’s Crissy Field salt marsh and sand dune restoration project (Jones and

Stokes 1996) recommended that no endangered species be included in the project,

primarily because project managers were concerned that some neighbors

expressed discomfort with the idea (N. Hornor, pers. comm. 1996). In San

Francisco, publicity over dog leash law enforcement for conservation of western

snowy plovers (Charadrius alexandrinus nivosus), a federally threatened species,

recently raised fears that endangered species would cause unwelcome and

burdensome increases in restrictions on recreational uses of urban parklands (D.

Hatch, pers. comm. 1994 to 1998, Miller et al. 1997, Golden Gate National

Recreation Area 1997). Similarly, removal of planted nonnative trees in San

Francisco for purposes of habitat restoration may engender strong opposition even

before restoration goals and methods are articulated (J. Sigg and P. Holloran,

pers. comm. 1998). Public outreach and education will be necessary to ensure

that unrealistic perceptions of regulatory burdens, recreational public land use,

and esthetic changes associated with habitat restoration and reintroduction do not
74

impede recovery actions for Raven’s manzanita. Schools, horticultural and

conservation organizations, and community organizations should be involved in

public outreach to prepare for implementation of recovery actions. Pilot projects

involving small-scale reintroduction of Raven’s manzanita, or authorized local

public horticultural displays (Reinartz 1995), should precede more ambitious

recovery actions to demonstrate and promote compatibility with urban land uses,

and to establish public confidence and cooperation.

5. Conservation Efforts

We proposed Raven’s manzanita for Federal listing as endangered on June 16,

1976. The species was listed as endangered on October 26, 1979 (U.S. Fish and

Wildlife Service 1979). It was also listed as endangered by the State of California

in 1978 (California Department of Fish and Game 1992). We published a final

recovery plan for the species on July 10, 1984, prepared by manzanita experts Jim

Roof and Alice Howard (U.S. Fish and Wildlife Service 1984). The recovery

priority number for Raven’s manzanita is 12, indicating a subspecies with a

moderate degree of threat and low recovery potential (see criteria published by

Federal Register Notice (48 FR 43098; September 21, 1983).

Since the publication of the 1984 recovery plan, the Golden Gate National

Recreation Area and its volunteers significantly reduced competition by

nonnative vegetation around the natural Raven’s manzanita clone, removing

Monterey cypress, iceplant, and nonnative grasses, and minimizing recolonization

by these species. Symbolic fencing, interpretive signs, and improved

coordination with road maintenance and other staff of the Golden Gate National

Recreation Area have reduced threats of trampling and accidental damage. These

actions have resulted in expansion of the clone in a generally healthy condition

most years. 

In January 1987, the Presidio (then managed by the U.S. Army) and the Golden

Gate National Recreation Area cooperatively propagated 168 cuttings of the wild

clone, from which 50 propagated plants survived to be planted in the Presidio. In

December 1987, the Golden Gate National Recreation Area established one small

colony of daughter clones around the parent clone, two small colonies in the

vicinity of the original plant, and one transplant about 1.6 kilometers (1mile)
75

away from it (U.S. Fish and Wildlife Service recovery file information). This

work is a step toward, but far short of, the 1984 recovery plan’s prescription for 5

populations of at least 20 plants each. The Golden Gate National Recreation Area

has prepared a comprehensive vegetation management plan for the Presidio

(Golden Gate National Recreation Area and Presidio Trust 2000, National Park

Service and Presidio Trust 2001), which proposes increased protection and

maintenance of the habitat of the mother and daughter clones at the World War II

Memorial site and transplants introduced to other Presidio locations. Tilden Park

(East Bay Regional Parks) and the University of California, Berkeley, have

maintained their collections of both endemic San Francisco manzanita taxa. 

Strybing Arboretum has maintained a single replicate clone of Raven’s

manzanita, but it has declined precariously in recent years due to blight and high

rainfall. The University of California, Berkeley, obtained open-pollinated and

self-pollinated seed from approximately 4,500 fruits harvested from cultivated

clones in 1995 (H. Forbes, pers. comm. 1999), from which 12 seedlings from the

open-pollinated source were obtained under experimental germination conditions;

it is unclear whether these seedlings are pure strains (selfed), hybrids, or mixed.

6. Species Recovery Strategy for Raven’s Manzanita

The recovery of Raven’s manzanita has three basic objectives, one of which has

been added to those prescribed in the original recovery plan (U.S. Fish and

Wildlife Service 1984). The first basic objective is to continue to protect the

existing remnant natural clone against foreseeable threats and ensure its natural

survival. This objective has been achieved, but requires ongoing effort. The

second objective is to reduce the chance of Raven’s manzanita’s extinction by

increasing the number of independent populations of the original clone at various

locations in the Presidio, spreading the risk of mortality among many replicate

colonies. These two objectives, continued from the original recovery plan, are

focused only on avoiding extinction of the species under highly managed

conditions, not on its continued evolution and conservation of the ecosystem on

which it depends (Endangered Species Act section 2(b), PL 100-478). Recovery

efforts under the original recovery plan were based primarily on protection of the

remnant clone, or on artificial propagation and transplanting of replicate clones;

there was limited emphasis on developing spontaneous reproduction of variable

plants in “self-sustaining” populations or habitat. This revised recovery plan now
76

includes objectives to restore sexual reproduction and regeneration of Raven’s

manzanita, and restore its potential natural ecological interactions with native

associated species, including potential rare gene flow between it and the related

Franciscan manzanita.

To make these multiple objectives compatible, the original remnant clone and its

associated native vegetation should be treated conservatively (protecting the local

plant assemblage in which it occurs, managing it by removing nonnative species). 

For the foreseeable future, additional Presidio populations should be clones of the

original, unless new “pure” clones (seedlings resulting from self-fertilization of

the original clone) are obtained. The Presidio should be a refuge for “pure”

Raven’s manzanita, either clones of the original or inbred seedlings, which will

require protection of existing suitable habitat or the establishment of new habitat. 

When Raven’s manzanita is reintroduced to existing suitable habitat, it is

important that new transplanted populations do not harm rare established native

serpentine vegetation. The most abundant opportunities for reintroduction occur

below Lincoln Boulevard on the serpentine bluffs (stable and unstable landslides),

where relatively bare serpentine soil slopes and serpentine bedrock outcrops are

still found. Exposed serpentine rocks and soils also occur behind Crissy Field,

near Fort Point, Inspiration Point, and a few other locations (McCarten 1986). 

Unexposed near-surface serpentine rocks and soils are also likely to occur along

the shear zone (Figure 3) elsewhere in the Presidio.

Multiple experimental populations of sexually reproductive Raven’s manzanita

should be established at interior San Francisco locations isolated from the

Presidio, preferably on serpentine outcrops near or similar to those of historic

(extirpated) localities. Because of the strong geographic and ecological isolation

between the Presidio and interior San Francisco localities, a wider range of

experimental reintroduction methods may be attempted with minimal risk to the

species. The establishment of new populations should be combined with

restoration of associated local serpentine plant species assemblages. Restoration

of appropriate plant assemblages would have to include reintroduction of

Franciscan manzanita in mixed populations. The relatively small size of such

urban reserves would be compatible with the historic natural distribution of

serpentine outcrop vegetation in San Francisco, which was a pattern of small,

discontinuous, localized sites.
77

No specific historic localities of Raven’s manzanita outside the Presidio are

known to contain potentially restorable serpentine habitat. However, potentially

restorable serpentinite-bearing sites occur at geologically related parts of the Fort

Point-Potrero Hill-Hunters Point shear zone (Figure 3). Potentially restorable

sites also occur within the same serpentinite-bearing formations elsewhere in San

Francisco along the City College shear zone (such as Lands End; Figure 3). 

Parks, undeveloped steep slopes, and road cuts in serpentine rocks represent

opportunities for both small-scale and large-scale reintroduction experiments. 

Examples of potential small urban reserves may be found near Potrero Hill (e.g.,

Starr King Park). Secure Federal lands with large serpentine outcrops occur

around the U.S. Mint at Duboce Street. Undeveloped land at the historic Mt.

Davidson locality, on greenstone, has potential for habitat restoration and

reintroduction. Other greenstone outcrops in undeveloped steep slopes occur

locally around Twin Peaks and Sunset Heights, and elsewhere (Figure 2; map in

Schlocker 1974). Other Franciscan rock outcrops with vegetation similar to that

of serpentine rocks could also be the subject of reintroduction experiments. 

Establishment of viable populations of Raven’s manzanita at these locations could

potentially provide habitat and populations that are roughly equivalent to those

that were extirpated.

Planning for experiments to reintroduce Raven’s manzanita into interior San

Francisco should begin by surveying contemporary conditions of mapped bedrock

outcrops of serpentine rocks and greenstone. Exposures should be assessed for

conservation potential (size, access, adjacent land use compatibility, maintenance

potential, vegetation restoration potential) and potential for management and

protection (public/private ownership, landowner interest, costs for easements or

fee title acquisition, compatibility with recreational or other land use, community

support) with serpentine being the higher priority. The San Francisco Department

of Parks and Recreation owns and manages many semi-wild vegetation remnants

in San Francisco, often in cooperation with local community groups, conservation

organizations, and schools.

The potential for sexual reproduction of offspring with adequate variability in

ecologically adaptive traits in Raven’s manzanita should be assessed. The highest

initial priority would be to generate new seedling-grown plants resulting from

self-fertilization (inbreeding) under controlled conditions. Viable seeds from
78

several hundred fruits were collected from cultivated plants at the University of

California, Berkeley. These seeds were reportedly produced by self-pollination

alone (H. Forbes, pers. comm. 1999). Currently, the University of California,

Berkeley Botanical Garden has 12 juvenile offspring derived from uncontrolled

open pollination of cultivated clones of Raven’s manzanita (H. Forbes, pers.

comm. 1999). These plants have not yet been cloned or tested for possible

hybridity. Additional attempts should be made to breed new “pure” genetic

individuals (“selfed” or inbred seedlings). The pedigrees of any plants resulting

from inbreeding plants would need to be recorded, and plants would need to be

genotyped (genetic individuals distinguished by heritable “marker” traits or

genes) and evaluated for indications of inbreeding depression (loss of fitness, i.e.

decreased viability or reproductive output, due to inbreeding). If the inbreeding

program yields significant numbers of viable plants, they should be propagated

clonally, and pedigreed clones should be reserved in cultivation, then

reintroduced to the wild or restored habitat.

Inbreeding may fail to generate viable cultivated stock of “pure” Raven’s

manzanita. Since the only seedlings successfully grown from Raven’s manzanita

fruits were from open-pollinated plants (no seedlings were obtained from viable

seed of selfed plants at the University of California, Berkeley; H. Forbes, pers.

comm. 1999), and no seedlings of Raven’s manzanita have ever been observed in

the wild, it is quite possible that this plant is an obligate outbreeder. Even if

viable inbred plants are produced, it is possible that the Presidio clone from the

windswept coastal bluff site may yield a limited range of genetic individuals that

would not be well adapted to the novel soil, vegetation, and microclimates of

interior San Francisco reintroduction sites. To restore levels of fertility and

genetic diversity to obligate outcrossing plant taxa that have been reduced to

single genetic individuals (e.g., Castilleja uliginosa, Physostegia correllii,

Hedyotis parvula, Betula murrayana, Pritchardia munroi, Prunus maritima var.

gravesii; McMahan 1989), one possible strategy is a breeding program based on

introgressive hybridization with closely related species, or genetically distinct

populations (Knapp and Connors 1999). Peter Raven and the late Robert Ornduff

had suggested applying this strategy to the sole survivor of the obligate

outcrossing species, Castilleja uliginosa (Falk 1992). Introgressive breeding is

part of the recovery plan for Kokia cookei, a Hawaiian plant that persists only as a

single genetic individual, in cultivation (U.S. Fish and Wildlife Service 1998d). 
79

Introgressive breeding is an extraordinary strategy for recovery of an extremely

rare plant, but ordinary concerns about the integrity of genetic diversity within

species may be effectively irrelevant when a species is reduced to a single

individual (Falk 1992). When the single survivor represents an obligate

outcrossing species that would be extremely unlikely to reproduce spontaneously,

extraordinary genetic management approaches may be appropriate (Falk 1992). 

Introgressive breeding is a strategy of last resort to recover near-extinct species

(Rieseberg 1991). It is based on developing first-generation (F1) hybrids between

the rare species (or distinct population) and a closely related species or

population, and crossing these F1 hybrids both back among themselves, and on the

rare parent species or population. Segregants of F2 hybrids (second generation

hybrids, individuals with various combinations of parental traits) would be

selected for those that are essentially indistinguishable from the rare parent. This

strategy would result in generations of individuals with genetic backgrounds

derived mostly from the rare species (depending on the number of backcrossed

generations), but with increased genetic diversity. This combination of

backcrossing and selection is a breeding strategy that is long established in

cultivated plants for horticultural and agricultural purposes (Briggs and Knowles

1965). For slow-growing woody species like manzanitas, this breeding program

would take many years or decades because of the time needed for each seedling

generation to reach sexual maturity. 

If attempts to generate genetically variable, viable, and fertile robust inbred lines

from the single Raven’s manzanita (the first priority) are unsuccessful, the

introgressive hybridization approach should be attempted as an alternative to

perpetual artificial propagation and planting to achieve survival of a single

asexual clone and as a last chance for continued evolution of the species. Before

attempting any introgressive hybridization efforts, however, evaluation of the

need and appropriateness for a breeding program for Raven’s manzanita should

be conducted by an expert scientific peer review panel. A genetic management

plan describing the breeding program and how any progeny would be used in

reintroduction should be developed collaboratively by us, the National Park

Service (Golden Gate National Recreation Area), the Presidio Trust, and the

scientific panel. The panel should include experts in manzanita biology, plant

conservation genetics, plant breeding, and plant propagation. 
80

A leading candidate taxon for an experimental introgressive breeding program

with Raven’s manzanita is Tamalpais manzanita (Arctostaphylos montana

[=Arctostaphylos hookeri ssp. montana]). It is geographically, ecologically,

morphologically, and taxonomically close to Raven’s manzanita (see Description

and Taxonomy). Recent genetic analysis (Markos et al. 1999) confirmed that this

species is related to its rare serpentine counterparts south of the Golden Gate, and

it shares the same number of chromosomes as Raven’s manzanita (Wells 1993). 

Wells (1993) treated both taxa as subspecies of the same species, and they are

distinguished by few morphological traits, some of which may be artifacts of lost

variability in Raven’s manzanita (Behr 1892, Roof 1976). Research is needed to

further clarify which manzanita taxa or populations would be most suitable for

inclusion in a breeding program with Raven’s manzanita, how compatible

genetically they are, and how the plants resulting from introgressive progeny

compare with Raven’s manzanita in ecological and morphological characteristics. 

Introgressive breeding of Raven’s manzanita introduces risks that intercross

individuals may mistakenly be planted as “pure” Raven’s manzanita (label error,

propagation errors). Such errors could potentially result in gene flow to the

“pristine” Presidio population or confusion between “pure” clones and intercross

individuals. If introgressive plants are bred, they should be restricted to isolated

interior San Francisco locations, and separated from the Presidio. However, the

concept of “pure” manzanita species is sometimes considered artificial (see

Description and Taxonomy) given the strong indications of widespread natural

hybridization in the evolution of the genus (Gottlieb 1968, Keeley 1976, Ellstrand

et al. 1987, Wells 1991, Schierenbeck et al. 1992).

Because of the vulnerability of the single existing plant of Raven’s manzanita,

conservation in botanical gardens will continue to play a role as a hedge against

extinction in the wild. Cultivation, however, is not an alternative to recovery in

wild populations. Cultivation provides benefits for scientific and educational

objectives, and for public outreach, but cannot achieve the Endangered Species

Act’s basic purpose of conserving endangered species in their natural ecosystems. 

Cultivated populations are essentially static and lack dynamic ecological and

evolutionary processes.
81

III. OTHER FEDERALLY LISTED SPECIES AND SPECIES OF

CONCERN OR REGIONAL CONSERVATION SIGNIFICANCE

The principal causes of decline and endangerment of San Francisco lessingia and

Raven’s manzanita have been loss of populations and habitat caused by

urbanization of San Francisco, and degradation of remnant habitat quality. The

habitat destruction associated with San Francisco’s urban growth has significantly

impoverished the flora of the San Francisco Peninsula, and also contributed

significantly to the decline of other federally listed species that are native there. 

These species include California sea-blite (Suaeda californica; U.S. Fish and

Wildlife Service 1994a), marsh sandwort (Arenaria paludicola; Behr 1892, U.S.

Fish and Wildlife Service 1998b), beach layia (Layia carnosa; U.S. Fish and

Wildlife Service 1998b), Marin dwarf-flax (Hesperolinon congestum; U.S. Fish

and Wildlife Service 1998a), and Presidio clarkia (Clarkia franciscana; U.S. Fish

and Wildlife Service 1998a). 

Urbanization also has caused or contributed to the decline of a host of other plant

species, including some that are now extirpated over parts of their range, nearly or

entirely extinct in the wild, or declining toward rarity and endangerment. These

species of concern are not currently federally listed as threatened or endangered, but

many could become so in the future. Some may be proposed for listing once adequate

survey information is available. Other species of local or regional conservation

significance include those that remain relatively secure in populations elsewhere, but

have undergone substantial range reduction, and have become rare or locally

extirpated on the San Francisco Peninsula.

The fundamental aim of this recovery plan is to address the conservation needs of its

endangered species within their ecosystems, including their associated species of

concern, and other species of regional conservation significance. Recovery actions

aimed at protection, management, and restoration of habitat and populations of listed

species should incorporate appropriate conservation actions for these associated

species.

Site-specific actions undertaken pursuant to this recovery plan may include

reintroduction, removal or eradication of nonnative invasive vegetation to indirectly

benefit the native plants, or temporary protective measures to avoid adverse impacts to
82

existing populations in the course of habitat restoration. Such actions must be

consistent with any existing recovery plans covering listed species on the Presidio.

A. Ecologically Associated Federal Listed Species

Several other federally listed species occur within the geographic and ecological scope

of this plan: Presidio clarkia (Clarkia franciscana), Marin dwarf-flax (Hesperolinon

congestum), and beach layia (Layia carnosa). Serpentine and sand dune habitats of

San Francisco are also within the historic range of the bay checkerspot butterfly

(Euphydryas editha bayensis) and Myrtle’s silverspot butterfly (Speyeria zerene

myrtleae). These species are covered comprehensively throughout their full

geographic ranges in two other recovery plans: the Recovery Plan for Serpentine Soil

Species of the San Francisco Bay Area (covering bay checkerspot butterfly, Presidio

clarkia, and Marin dwarf-flax; U.S. Fish and Wildlife Service 1998a) and the

Recovery Plan for Seven Coastal Plants and the Myrtle’s Silverspot Butterfly

(covering Myrtle’s silverspot butterfly and beach layia; U.S. Fish and Wildlife Service

1998b). The reader is referred to these documents for more extensive treatment of the

recovery needs of these species. Below are summary accounts of these species. 

Recovery actions and recommendations from all relevant recovery plans should be

incorporated in areas where these species are ecologically associated with the species

covered in this recovery plan. For example, Presidio clarkia was seeded into the

serpentine habitat of Raven’s manzanita nearly three decades ago (Roof 1972), even

before its Federal listing, and has persisted since then.

1. Presidio Clarkia, Clarkia franciscana Harlan Lewis & Raven 

Presidio clarkia (Clarkia franciscana) is a slender, erect annual herb in the eveningprimrose family (Onagraceae) (Figure 9). The ecology and recovery of this species

are covered comprehensively in the Recovery Plan for Serpentine Soil Species of the

San Francisco Bay Area (U.S. Fish and Wildlife Service 1998a). It has showy pink

and purple flowers in spring and early summer on grasslands of serpentine soils and

outcrops. Presidio clarkia is similar to Clarkia rubicunda, which occurs in open

grassland and scrub vegetation on the San Francisco Peninsula (McClintock et al.

1990), but Clarkia rubicunda is not a close ancestor of Presidio clarkia, as had

previously been concluded (Gottlieb 1973). Presidio clarkia is a predominantly selfpollinating plant (Lewis and Raven 1958). Population sizes fluctuate strongly among

years. In some years populations drop to zero (Roof 1971), and apparently regenerate
83

Figure 9. Selected plant species from the northern San Francisco Peninsula

that are ecologically associated with either serpentine or dune

vegetation. Species are ordered alphabetically by their botanical

name. 

INDIAN PAINTBRUSH

Castilleja affinis ssp. affinis

NUTTALL’S MILK-VETCH

Astragalus nuttallii var. virgatus

PINK SAND-VERBENA

Abronia umbellata ssp. umbellata

FRANCISCAN MANZANITA

Arctostaphylos franciscana

(=A. hookeri ssp. franciscana )
84

Figure 9 (continued). Selected plant species from the northern San Francisco

Peninsula.

DUNE TANSY

Tanacetum camphoratum

SEASHORE STARWORT

Stellaria littoralis
85

CURLY-LEAVED MONARDELLA BEACH LAYIA

Monardella undulata Layia carnosa

(federally listed as endangered)

 COMMON LINANTHUS COAST PIPERIA

 Linanthus parviflorus Piperia elegans

Figure 9 (continued). Selected plant species from the northern San Francisco

Peninsula.
86

 SAN FRANCISCO SPINEFLOWER

 Chorizanthe cuspidata

BANDED PURPLE OWL’S-CLOVER

 Castilleja exserta ssp. latifolia

 FRANCISCAN THISTLE PRESIDIO 

Cirsium andrewsii CLARKIA

 Clarkia franciscana

 (federally listed

 as endangered)

Figure 9 (continued). Selected plant species from the northern San Francisco

Peninsula. 
87

SAN FRANCISCO DUNE GILIA

 Gilia capitata ssp. chamissonis

FRANCISCAN WALLFLOWER

 Erysimum franciscanum

 SAN FRANCISCO GUMPLANT

 Grindelia hirsutula ssp. maritima

MARIN DWARF-FLAX 

Hesperolinon congestum

 (federally listed as threatened)

Figure 9 (continued). Selected plant species from the northern San Francisco

Peninsula.
88

from seedbanks in more favorable years like other native annuals. Experience with

vegetation management in the Presidio suggests that Presidio clarkia’s reproductive

success may depend on the competing vegetation being sparse. The remnant wild

Presidio population was established on a former serpentine rock quarry (Roof 1971).

Population declines appear to be associated with encroachment by nonnative

vegetation, particularly dense growth of annual nonnative grasses (M. Albert and S.

Farrell, pers. comm. 1998). 

Presidio clarkia is known from only two natural populations: one in San Francisco

(Inspiration Point, Presidio) and a series of subpopulations in the Oakland Hills

(Gottlieb and Edwards 1992). An artificially seeded population in the Presidio (World

War II Memorial, adjacent to the parent clone of Raven’s manzanita) occurs on a

coastal serpentine outcrop above the north end of Baker Beach. It was established

there by translocation of seed in 1972 (Roof 1972), and has persisted since then at

population sizes ranging from tens to hundreds of plants. In 1994, 860 plants

established at this introduced population, while 8,716 grew at the managed natural

Inspiration Point population (Golden Gate National Recreation Area unpublished data

1994). The main population on the Presidio is protected against development, but it

remains strongly threatened by introduced conifers and blue gum (Eucalyptus

globulus) trees, weedy herbaceous nonnative plant species, trampling, and unfavorable

mowing times (prior to seed maturation and dispersal). Potential development of

Presidio Trust lands currently unoccupied by Presidio clarkia, but containing

serpentine outcrops and soils, may preempt habitat restoration that may be needed for

recovery of this species. Because some recovery actions for Raven’s manzanita in

serpentine habitat within the Presidio may benefit Presidio clarkia within manzanita

habitat (as demonstrated by the successful establishment of the World War II

Memorial population of Presidio clarkia) recovery actions for Presidio clarkia should

be integrated with those of Raven’s manzanita. Recovery actions for Raven’s

manzanita that may benefit Presidio clarkia include enhancement of serpentine habitat

quality (e.g., removal and suppression of invasive nonnative grasses, iceplant

(Carpobrotus edulis), and conifers). Resources for recovery of both endangered

species could be allocated efficiently by reintroducing both to unoccupied suitable

serpentine outcrop habitat on the Presidio bluffs. 
89

2. Marin Dwarf-flax, Hesperolinon congestum (Gray) Small

Marin dwarf-flax (Hesperolinon congestum) is a small (usually less than 15

centimeters [6 inches]) annual herb in the flax family (Linaceae) (Figure 9). This

federally threatened species is addressed in the Recovery Plan for Serpentine Soil

Species of the San Francisco Bay Area (U.S. Fish and Wildlife Service 1998a). It has

threadlike stems and linear leaves. Its flowers, which have five rose-to-whitish petals

and deep pink or purple anthers, are borne in clusters. Marin dwarf-flax is known

from fewer than 20 serpentine soil localities in Marin, San Francisco, and San Mateo

Counties. It is closely related to the more widespread California dwarf-flax

(Hesperolinon californicum), from which it is distinguished by its hairy sepals, dark

pink/purple anthers, and exclusive serpentine habitat. It occurs in serpentine grassland

soil outcrops above Baker Beach, near the one remaining natural Raven’s manzanita

location. California dwarf-flax, but not Marin dwarf-flax, co-occurs with Raven’s

manzanita at the World War II Memorial site. Marin dwarf-flax was historically

reported from the former Laurel Hill Cemetery (now urbanized) and Lone Mountain

(urbanized) around the turn of the 20th century (Howell et al. 1958). Its decline in

San Francisco is attributable to urban development, invasion by dominant nonnative

vegetation, and trampling. 

3. Beach Layia, Layia carnosa (Nuttall) Torrey and Gray

Beach layia (Layia carnosa) is a prostrate to low erect annual fleshy-leaved herb in the

aster family (Asteraceae) (Figure 9). The biology and ecology of this plant are

summarized in the Recovery Plan for Seven Coastal Plants and Myrtle’s Silverspot

Butterfly (U.S. Fish and Wildlife Service 1998a). Beach layia germinates during

winter rains and grows as a vegetative rosette until spring, when flowering shoots bolt. 

Individual plants produce from 1 to over 100 seed heads. Seeds (achenes) have a

bristly pappus that facilitates wind dispersal. The species is locally common, but

occurs in few populations that are widely scattered from Humboldt County to Santa

Barbara. It occurs only in sparsely vegetated coastal dunes in the Humboldt Bay area,

Point Reyes, Monterey Peninsula, and Vandenberg Air Force Base. The largest

populations occur in dunes around Humboldt Bay and Point Reyes.

Beach layia was reported from San Francisco (Thomas 1961), although no herbarium

specimens from San Francisco are found in the herbaria of the California Academy of
90

Sciences, Dudley/Stanford (P. Baye, pers. observ. 1997), Jepson, or University of

California (CALFLORA 1999). Unless it was misreported, it probably consisted of

small local populations in San Francisco, since it was not detected by the preparers of

San Francisco floras prior to 1961 (Brandegee 1892, Howell et al. 1958). It is

currently absent in San Francisco, where suitable habitat for reintroduction exists (e.g.,

portions of Ocean Beach, Golden Gate National Recreation Area; Crissy Field dune

restoration area; and Fort Funston dunes), and can be expanded. Reintroduction and

habitat expansion would be consistent with the approved recovery plan for this

species.

Beach layia occurs in a wide range of successional phases of coastal foredunes, dune

grassland, and scrub, but is always associated with sparse, open sandy areas. At Point

Reyes, the nearest population to San Francisco, it occurs in semi-stable foredunes,

sparsely vegetated blowouts in dune scrub, and active bare dunes. It grows in Point

Reyes dunes in both relatively recently deposited dune sand as well as in openings in

older dunes with relatively weathered, organically enriched old sandy soil (comparable

to modern localities of San Francisco lessingia) (P. Baye, pers. observ. 1985 to 1999). 

Reproductive output of beach layia corresponds with plant size, and both are relatively

greater in microenvironments with elevated soil moisture (edges of dune slacks and

lagoons) or organic matter (older dune soils). Population size of this annual species

fluctuates strongly from year to year. It is endangered by loss of habitat and

degradation of its habitat due to extensive establishment of European beachgrass

(Ammophila arenaria), as well as iceplant and other invasive nonnative vegetation

(U.S. Fish and Wildlife Service 1998b). 

4. Myrtle’s Silverspot Butterfly, Speyeria zerene myrtleae 

Myrtle’s silverspot butterfly (Speyeria zerene myrtleae), an endangered member of the

brush-footed butterfly family (Nymphalidae), historically ranged from coastal San

Mateo County (Pescadero) to northern Sonoma County. It inhabits coastal dunes,

coastal scrub, and coastal grasslands that support adequate nectar sources and the

larval host plant, western dog violet (Viola adunca) (U.S. Fish and Wildlife Service

1998b). The host plant occurred historically in grassland at localities of both historic

and remnant dunes in San Francisco (Baker Beach, Sunset Heights; Howell et al.

1958). Though now rare or locally extirpated at these and other localities of existing
91

and proposed San Francisco lessingia populations, dog violet and possibly Myrtle’s

silverspot butterfly could potentially recolonize or be reintroduced to San Francisco

lessingia recovery units. Such reestablishment would be consistent with the approved

recovery plan for this species.

5. Bay Checkerspot Butterfly, Euphydryas editha bayensis 

The threatened bay checkerspot butterfly is associated with serpentine grassland

habitats, but it also occurred historically at Twin Peaks in San Francisco (not a

serpentine area, but bearing other Franciscan bedrock outcrops) and San Bruno

Mountain (U.S. Fish and Wildlife Service 1998b). Its primary larval food plant is

dwarf plantain (Plantago erecta; U.S. Fish and Wildlife Service 1998b). Dwarf

plantain is still locally abundant in San Francisco and San Mateo County, occurring on

remnant and restored older sand dunes with some soil development, and on thin soils

of Franciscan bedrock outcrops composed of greenstone, serpentine, or sandy

Pleistocene sediments (P. Baye, pers. observ. 1996 to 1999). Secondary, alternative

larval host plants include purple owl’s-clover (Castilleja exserta = Orthocarpus

purpurascens), a species of local conservation significance (see section C below) that

is extirpated in San Francisco (P. Baye, pers. observ. 1984 to 1999), but persists

nearby on coastal grasslands at headlands in Pacifica, San Mateo County (Rockaway

Head; P. Baye, pers. observ. 1999) and locally in older Point Reyes dunes (P. Baye,

pers. observ. 1999). Purple owl’s-clover was formerly locally abundant in dunes and

coastal grasslands of San Francisco (Howell et al. 1958).

Restoration of coastal dune grassland and Franciscan bedrock outcrops (particularly

where they occur together in large potential reserve areas - e.g., Sunset Heights, Baker

Beach), with reintroduction of the host plant species, would provide opportunities for

reintroduction or spontaneous recolonization of a bay checkerspot butterfly

population. Such reestablishment would be consistent with the approved recovery

plan for this species.

B. Species of Concern 

The following species occurred in the historic flora of San Francisco and northern San

Mateo County as components of dune or bedrock outcrop vegetation, and are either

known or probable associates of Raven’s manzanita or San Francisco lessingia. They
92

are now either in major local decline, locally rare, locally extirpated, or locally extinct

in the wild (surviving only in cultivation). Some of these rare species have few or no

recent records or survey information, and may warrant future listing as threatened or

endangered species. The plant assemblages in which they were included were part of

the ecological and evolutionary background of the two endangered species featured in

this plan. Recovery of the ecosystem in which these listed species occur should

include restoration of the approximate vegetation and flora it supported. 

Little is known about the interactions between San Francisco lessingia or Raven’s

manzanita and their native plant associates. Some interactions may be antagonistic,

such as shading by taller shrubs or nutrient or water competition below ground. But

interactions between plants may be positive at some life-history stages, such as partial

foliar shading, which reduces moisture stress of seedlings, or persistent leaf litter,

which traps seeds in favorable microsites. Some native competitors may help exclude

establishment of nonnative invasive species that are more antagonistic to listed

species. Other potential interactions include attraction of insect pollinators,

modification of soil nitrogen content (nitrogen fixation), extraction of nutrients from

different strata of the soil profile, and sheltering or “nurse plant” effects on seedlings. 

The relative frequency of contacts among less common species is also unknown or

poorly known; some may have directly associated with one another, and some may

have occupied distinctly different microenvironmental patches within the same

vegetation. It is impractical (or impossible) to perform an adequate comprehensive

study of plant interactions in local dune and rock outcrop vegetation. Thus, it is

prudent to reassemble the historic elements of the vegetation to the extent possible,

based on historic records and assessment of representative stands of similar vegetation

in the region, and allow community dynamics to reconstitute themselves to some

degree. The reconstitution of serpentine and dune plant communities dominated by

the native species that were historically present is essential to full recovery of Raven’s

manzanita and San Francisco lessingia.

The most basic tenet of restoration of plant communities is not to lose the parts that

compose them. Some of the plant species of concern have apparently been lost in the

San Francisco flora, at least outside of cultivation. Some species persist in San

Francisco, but have been extirpated in distinct types of plant communities, such as

dune scrub and grassland, where local, ecologically distinctive populations are likely

to evolve. It also is possible that some apparently extirpated species are “latently”
93

present as dormant seeds in the soil, awaiting ecological regeneration following

disturbances to soil and vegetation (Thompson 1992). Conservation of these species

should rely first on protecting and, where feasible, expanding existing remnant

populations, particularly in dune and rock outcrop habitats. The first priority for

conservation of species of concern should be to expand local populations rather than

import propagules from offsite, to retain the character of local populations. 

Populations can sometimes be expanded by enhancing habitat quality and allowing

natural spread, which is a preferable method. In remnant natural areas of San

Francisco, population expansion is often made possible by removal of nonnative

plants, thwarting their re-invasion, and managing disturbances such as excessive

trampling. Where conditions for establishment are more difficult, artificial

propagation of local stock or seed and outplanting (population augmentation rather

than reintroduction) may be appropriate.

For species known or presumed to be extirpated, systematic surveys should be

conducted in suitable habitat at appropriate seasons for detection. For species that are

locally extirpated, reintroduction from appropriate source populations should be

attempted experimentally. Reintroductions of these species of concern to places where

they have been extirpated should be presumed to be difficult and will have to be

conducted on an experimental basis because of the many uncontrolled variables

affecting restored habitats (rainfall and drought, pathogens, herbivory, nonnative

competitors). Like natural dispersal and establishment of founder populations

(Primack 1996), reintroductions often have low probabilities of success, and may

require much repetition under variable conditions to succeed (Falk 1992). They are

also uncertain because little is usually known of the biology of the species being

reintroduced, such as genetic population structure, pollinators, and requirements for

germination or seedling establishment. Therefore, reintroductions are likely to require

“adaptive management” tactics in horticultural and ecological management practices,

and much experimental replication (trial and error). Reintroduction may also depend

on developing sound practical understanding of horticultural considerations

(propagation, transplanting techniques, soil preparation, seasonal timing; Guerrant

1996) and ecology (local variation in microhabitat preferences, adult and seedling

habitat, pollinators, pathogens, herbivory) for individual species. Important local

ecological information may be essential for successful reintroduction, but is seldom

available in the biological literature. 
2

 Hemiparasitic: a plant that parasitizes other plants by attaching to their roots, but

nevertheless has photosynthetic leaves of its own. 

94

Not all plant species reintroductions are difficult and failure-prone, however. Some

reintroduced native species appear to be naturally robust colonizers of earlysuccession habitats, and sometimes establish readily in suitable conditions. Many

native coastal dune species such as sea-rye or dunegrass (Leymus mollis), sand

verbenas (Abronia latifolia and the uncommon introgressant Abronia umbellata),

beach strawberry (Fragaria chiloensis), coyote-brush (Baccharis pilularis), and beach

evening-primrose (Camissonia cheiranthifolia) fit this category. They spontaneously

colonize restoration sites as well as degraded habitats. Many other native shrubs,

grasses, and creeping herbs, both widespread and uncommon taxa, can be robust

colonizers of suitable habitat. Uncommon hemiparasitic2

 paintbrushes (Castilleja

wightii and Castilleja affinis ssp. affinis), which are reputedly difficult to cultivate,

spontaneously colonize iceplant-covered roadway medians and roadsides and road

cuts from San Francisco to Montara, San Mateo County (P. Baye, pers. observ. 1998

to 1999). Even the endangered San Francisco lessingia shows opportunistic “weedy”

tendencies in disturbed sandy areas.

Of particular concern is the provenance of stock populations for reintroduction

(Guerrant 1992, 1996; Knapp and Rice 1994; Lesica and Allendorf 1999). If local

populations are confirmed to be extirpated, or are severely depleted, supplemental

populations should be sought from closely similar ecological conditions (soil type,

aspect, exposure) from the closest populations, in view of relevant mechanisms and

pathways of dispersal. If source populations are themselves extremely depleted in

numbers (and presumably, though not necessarily, genetic diversity), combining

mixed founders from different suitable source populations should be evaluated (Lesica

and Allendorf 1999). Each species would require careful individual evaluation for

optimal reintroduction strategies, including surveys for remnant local and potential

offsite source populations, and propagation or transplanting techniques. These

assessments and plans should be implemented in restoration and reintroduction

projects aimed at listed species. 
95

1. Franciscan Manzanita, Arctostaphylos franciscana Eastwood 

 

Franciscan manzanita (Arctostaphylos hookeri G. Don ssp. franciscana (Eastwood,

Munz) is a low, spreading to ascending evergreen shrub in the heath family

(Ericaceae) (Figure 9). It is extinct in the wild, but formerly occurred mixed with

Raven’s manzanita on serpentine outcrops at three of four historically recorded

localities. It is the diploid (n = 13) ecological equivalent of the tetraploid (n = 26)

Raven’s manzanita. Recent molecular genetic data suggest that it is possibly a parent

species, or a derivative of a parent species, of Raven’s manzanita (M. Vasey, pers.

comm. 1998; Markos et al. 1999). These data also suggest that Raven’s and

Franciscan manzanitas are not closely related to A. hookeri subspecies of the Monterey

coast, but probably constitute a distinct species complex also including Tamalpais

manzanita (Arctostaphylos montana Eastwood, treated as Arctostaphylos hookeri ssp.

montana by Wells [1993]). Franciscan manzanita was once a candidate for Federal

listing (U.S. Fish and Wildlife Service 1980) (45 FR 82480), but was not listed

because no natural populations existed in the wild. 

Mixed wild populations of Franciscan and Raven’s manzanita occurred at the Laurel

Hill Cemetery, the Masonic Cemetery, Mt. Davidson (Howell et al. 1958, U.S. Fish

and Wildlife Service 1984), and possibly the Haight/Laguna Street site reported by

Behr (1892) as well. At Laurel Hill, Franciscan manzanita was allegedly less

abundant than Raven’s manzanita (Roof 1976), where both associated with coast live

oak (Quercus agrifolia), blue-blossom (Ceanothus thyrsiflorus), and coyote brush

(Baccharis pilularis), according to Wieslander’s herbarium collection (Wieslander

838, 27 October 1938). A collection label by L. Edmonds from the same locality

(February 15, 1958) noted that it grew “in leafmold on serpentine outcrop.” Behr

(1892) referred to the 1850's occurrence of the “tall form of Manzanita still growing so

abundantly on the slopes of Tamalpais” (Arctostaphylos montana Eastwood,

considered also by Markos et al. [1999] to be closely related to Raven’s manzanita) at

the site of the Protestant Orphan Asylum. Brandegee (1892) cited the manzanita at

this site as “Arctostaphylos pungens HBK,” located at Laguna and Haight streets. The

San Francisco “Arctostaphylos pungens” of these authors is interpreted as either

Franciscan manzanita, or mixed populations of this species and Raven’s manzanita, in

sheltered conditions that promote the growth of tall, branched plants (U.S. Fish and

Wildlife Service 1984). Like Raven’s manzanita, Franciscan manzanita is endemic to
96

bedrock outcrops (primarily serpentine, but including greenstone, and likely mixed

Franciscan rocks) in San Francisco. 

While Franciscan manzanita has been extinct in the wild since the mid-20th century,

up to three apparently distinct clones are maintained in cultivation. Differences

between clones persist after propagation, suggesting that they are genetic individuals

(D. Mahoney, pers. comm. 1998). Roof (1980) reported that he salvaged plants of

Franciscan manzanita from the Laurel Hill site between 1938 and 1940, possibly

multiple clones, from which modern collections are derived. These cultivated plants

vary in degrees of prostrate and ascending growth habits, and are less susceptible to

twig blight than Raven’s manzanita in years of high rainfall (P. Baye, pers. observ.

1997 to 1999). The plants are propagated by vegetative cuttings with moderate

difficulty but are easily cultivated, and thrive on neglect after they are established on a

wide range of substrates, including dune sand in Strybing Arboretum, not a natural

substrate for this shrub. Successful growth on substrates markedly different from

serpentine suggests good soil adaptability and a high potential for reintroduction on

various substrates. Franciscan manzanita also sets viable seed (possibly hybrid) that

can be propagated as well, reportedly forming highly variable progeny (D. Mahoney,

pers. comm. 1998). 

The historic association between the two endemic San Francisco manzanitas at most

interior historic San Francisco localities (not the Presidio, which is outside the known

range of Franciscan manzanita) suggests that they should be reintroduced together at

new (or formerly occupied) sites to allow for the possibility of rare hybrids and

interspecific gene flow in the continued evolution of the complex, and to allow for

other ecological interactions between these similar species. Reintroduced plants

should be propagated only from either clones or seed from controlled pollination of

inbred lines of Franciscan manzanita, if feasible. Reintroduction may be attempted on

both serpentine substrates and other types of Franciscan rocks.

2. Franciscan Thistle, Cirsium andrewsii Gray

Franciscan thistle is a coarse, spiny, short-lived perennial herb in the aster family

(Asteraceae), growing from 0.6 to 2.0 meters (2 to 6 feet) tall when mature (Figure 9). 

It grows from a large low rosette when juvenile. Franciscan thistle is rare, found in

small and often isolated populations on wet soils and seeps of coastal bluffs, streams
97

and canyons (Munz 1959, Keil and Turner 1993). At the Presidio, populations of

Franciscan thistle appear to have an affinity for vegetation gaps in both undisturbed

and disturbed soils and successional habitats within low-growing coastal scrub (P.

Baye, pers. observ. 1998). Some populations of this species in the Marin Headlands

appear to regenerate in relatively closed wet coastal scrub vegetation. Hybrids with

brownie thistle (Cirsium quercetorum) have been reported from Point Reyes (Howell

1949).

Franciscan thistle is reported to range from northern San Mateo (Thomas 1961) to

Sonoma Counties (Munz 1959), but no collections from Sonoma County were

reported in the flora of Sonoma County (Best et al. 1996), and it is not reported in the

flora of San Bruno Mountain (McClintock et al. 1990), which includes the mostly

likely remaining habitat in San Mateo County. Only a few historic and current

populations in Marin County have been reported: Rodeo Lagoon (apparently

extirpated), Tennessee Cove/Valley (confirmed 1985 by Golden Gate National

Recreation Area staff), Gerbode Valley (reconfirmed by Golden Gate National

Recreation Area staff early 1999), Dillon Beach, and Point Reyes. San Francisco

populations have been recorded only by J. Howell in the Presidio and near Lake

Merced. Franciscan thistle was historically collected east of Lake Merced in wet

ground near a willow grove, and in serpentine seeps at Fort Point, Presidio (Howell et

al. 1958). Three colonies occur between Fort Point (on serpentine bluffs immediately

east of the Golden Gate Bridge) and Battery Boutelle, all on the marine bluffs. The

Fort Point colony is in decline (S. Farrell, pers. comm. 1999). A small but expanding

population, located in 1998 (at least five individuals large enough to flower, two of

which flowered in 1999; P. Baye unpublished data 1998 to 1999), is established and

managed near and in a mudslide on serpentine bluff seeps below Fort Scott (Battery

Boutelle). Another colony of about five plants was located in late summer of 1999

(M. Chasse and P. Brastow, pers. comm. 1999). The distribution and abundance of

Franciscan thistle needs reevaluation. This plant may be considerably rarer than has

been presumed based on historic localities. Many of its historic localities have not

been surveyed recently and may be extirpated. Surveys are particularly needed in San

Mateo County.
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3. San Francisco Spineflower, Chorizanthe cuspidata Watson

 San Francisco spineflower is a prostrate annual in the buckwheat family

(Polygonaceae) (Figure 9). It bears flowers and fruits with persistent spiny bracts that

readily attach to fur and fabric, facilitating seed dispersal. San Francisco spineflower

is ecologically restricted to sparse, open vegetation on sand or sandy soils. Taxonomic

treatment of this species, and its delimitation from closely related species, has changed

over decades, and is not consistent in variously dated floras (Howell 1949, Thomas

1961, Hickman 1993). Nearly all populations occur on coastal sand dunes; a few

occur on weakly consolidated sandstone. Within its narrow geographic range along

the immediate coast from San Mateo County to southern Sonoma County, it occurs in

local abundance in coastal sands of Bodega Head, Dillon Beach, Point Reyes, and

southwestern portions of the Presidio in San Francisco (Hickman 1993, Best et al.

1996, P. Baye unpublished data 1992 to 1999). It has been reported historically from

near Santa Cruz (Thomas 1961). In San Francisco it has been recorded at Baker

Beach, near the Cliff House, Richmond District, Lake Merced (Fort Funston), and

Presidio (Howell et al. 1958). Because of the dispersal and colonization ability of this

species, it is expected to have relatively dynamic local population distribution. 

Brandegee (1892) described it as widespread in the western part of the City. The type

locality is San Francisco. It is recognized as a species of concern in the Recovery Plan

for Seven Coastal Plants and the Myrtle’s Silverspot Butterfly (U.S. Fish and Wildlife

Service 1998b).

4. Round-headed Chinese Houses, Collinsia corymbosa Buist ex Graham 

This showy annual member of the snapdragon family (Scrophulariaceae) was collected

in San Francisco in the late 19th and early 20th century. According to Wood (1996) it

was collected at Mountain Lake (1891), Lake Merced (1900), and Ocean View (1892),

which are all general sandy localities known to have supported San Francisco

lessingia. Collections at the California Academy of Sciences include Presidio golf

links (former dunes) and unspecified localities within San Francisco. Round-headed

Chinese houses was historically reported from Bolinas Lagoon, and was collected by

A. Eastwood at Point Reyes (Howell 1949), but has not been confirmed in recent

decades. The species is not reported from Sonoma County according to Best et al.

(1996), but Skinner and Pavlik (1994) identify a need for verification of an historic
99

“Russian colony” collection (possibly either at Fort Ross, Sonoma County, or obtained

from then-unnamed coastal localities by collectors stationed there). 

Munz (1959) described round-headed Chinese houses as restricted to sandy coastal

environments from Humboldt to San Francisco Counties. It is related to, and may

intergrade with, the similar white Chinese-houses (Collinsia bartsiifolia [spelled

C. bartsiaefolia in the older literature]) (Howell et al. 1958). Jepson (1925) reported it

only from the central Mendocino coast (presumably Tenmile Dunes). It is presumed

to be extirpated in San Francisco. The only population that has recently been verified

is in dunes near Fort Bragg to the Tenmile River, Mendocino County; this population

is possibly the only one remaining anywhere. Other historic localities in Humboldt

County urgently need verification. This species is threatened by incipient invasion by

European beachgrass at Tenmile Dunes, a weed that could permanently eliminate its

habitat if allowed to spread (U.S. Fish and Wildlife Service 1998b). Round-headed

Chinese houses is also vulnerable to random extinction because of small population

size and extremely narrow geographic range.

5. San Francisco Wallflower, Erysimum franciscanum Rossbach 

San Francisco wallflower is a short-lived subshrubby perennial herb in the mustard

family (Brassicaceae) with showy, sweet-scented cream to cream-yellow flowers

(Figure 9). In San Francisco, it is ecologically wide-ranging in relatively open, low, or

sparse ground layer vegetation of rock outcrops, serpentine soils, and dunes (P. Baye,

pers. observ. 1990's). San Francisco wallflower was recorded historically in San

Francisco at Baker Beach, Point Lobos (1912), Lone Mountain, Sunset Heights,

Laguna Honda (1933), and Lake Merced (1901) (Howell et al. 1958, Wood 1996). 

Remnant populations occur at the Sunset Heights Dune remnants and Franciscan

outcrops, Baker Beach, and Fort Funston (Golden Gate National Recreation Area

unpublished data; P. Baye, pers. observ. 1998). Nearby scattered populations occur on

San Bruno Mountain (McClintock et al. 1990), along Highway 1 from Pacifica and

Montara, and in the Marin Headlands (P. Baye, pers. observ. 1990's). It is rare in

Marin, Santa Clara, Santa Cruz, San Mateo, and Sonoma Counties (Skinner and Pavlik

1994).
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6. San Francisco Gumplant, Grindelia hirsutula Hook. & Arn. var. maritima (E.

Greene) M. A. Lane

San Francisco gumplant is a prostrate subshrub of the aster family (Asteraceae)

(Figure 9). It occurs on serpentine and sandstone coastal bluffs, and sometimes on

rock outcrops away from the immediate coast, where vegetative cover is sparse and

open. San Francisco gumplant is variable in its morphological traits, and many

populations are somewhat ambiguous taxonomically (Lane 1993). Another prostrate

gumplant that grows on coastal bluffs and rocks within San Francisco gumplant’s

range is coastal gumplant (Grindelia stricta ssp. platyphylla), which has broader,

rounded leaves and pappus awns (small barb-like appendages on seed-like dry fruits)

that are less than 0.3 millimeters (0.01 inch) long and lack teeth (Lane 1993).

San Francisco gumplant is uncommon to rare in Monterey, Marin, Santa Cruz, San

Luis Obispo, and San Mateo Counties; current localities need verification (Skinner

and Pavlik 1994). In San Francisco it appears to occur relatively early in succession

after episodes of bluff erosion (P. Baye, pers. observ. 1990's). In San Francisco, it was

historically recorded on Presidio bluffs, Point Lobos, Lake Merced, Laguna Honda,

and Twin Peaks (Howell et al. 1958, Wood 1996). A local form with hairy stems

(formerly recognized as Grindelia maritima (E. Greene) Steyermark forma anomala

Steyermark) co-occurred with typical Grindelia hirsutula var. maritima at Laguna

Honda, Twin Peaks, and San Miguel Hills (Howell et al. 1958). Currently it occurs

with Raven’s manzanita (partly due to planting), and numerous colonies occur along

the serpentine bluff area of the Presidio. Apparent hybrids occur in northern San

Mateo County, according to McClintock et al. (1990).

7. Diablo Helianthella, Helianthella castanea E. Greene

Diablo helianthella is a taprooted sunflower-like perennial herb in the aster family

(Asteraceae). It grows from a caudex (swollen trunk-like base bearing multiple

herbaceous shoots). It resembles common mule-ears (Wyethia spp.) in general

appearance. The species is rare, occurring in open grassland on interior hills of the

east Bay and San Francisco Peninsula. It is extirpated in San Francisco (Wood 1996)

where it formerly occurred near Ocean View, southeastern San Francisco Hills and

Bayview Hills in the late 19th century (Brandegee 1892, Howell et al. 1958). It is not

known which specific local plant associations included Diablo helianthella, but
101

grasslands and sparse scrub on thin, rocky soils (such as bedrock outcrop slopes) are

probable candidates. The nearest reported population occurs on San Bruno Mountain

in Brisbane (McClintock et al. 1990).

8. Evax, Hesperevax sparsiflora (A. Gray) E. Greene

Evax is a short (less than 15 centimeters [5 inches]) woolly-leaved annual herb in the

aster family (Asteraceae) with multiple ascending to erect stems. On the San

Francisco Peninsula and the adjacent region, evax occurs on rocky, thin soils and

grasslands (Thomas 1961). In San Francisco, it occurred primarily on serpentine soils

(Howell et al. 1958). The treatment in the Jepson Manual (Hickman 1993) by

Morefield (1993) divides the species into two varieties: plants from serpentine are

Hesperevax sparsiflora var. sparsiflora while plants from sandy bluffs and flats are

designated as Hesperevax sparsiflora var. brevifolia, which is described as

“uncommon.” The two varieties of evax are separated by characters with overlapping

variation (leaf length, relative leaf roundness, hairs), so there may be some question

regarding the varietal identity of the San Francisco plants. Previously, San Francisco

plants had been recognized by Howell et al. (1958) as Evax sparsiflora (A. Gray)

Jepson without varietal distinctions. The historical distribution of the San Francisco

plants is from the Presidio, Potrero Hills, Hunters Point, and McLaren Park (Howell et

al. 1958, Wood 1996) – all localities including serpentine outcrops, and all but the

Presidio lacking sandy substrates, so one would expect these plants to be assigned to

Hesperevax sparsiflora var. sparsiflora. Wood (1996) instead assigned the San

Francisco plants to Hesperevax sparsiflora var. brevifolia. 

The contemporary local abundance of evax on the San Francisco Peninsula is

unknown, but it may now be locally extirpated. Additional survey information on this

species is needed in this region.

9. Kellogg’s Horkelia, Horkelia cuneata Lindley ssp. sericea (A. Gray) Keck

Kellogg’s horkelia is a resinous-scented perennial herb in the rose family (Rosaceae)

that grows from a branched semi-woody base. Its leaves are conspicuously covered

with silky pubescence, and glands are inconspicuous or absent. This and similar

species (Marin horkelia [Horkelia marinensis] and wedge-leaf horkelia [Horkelia

cuneata ssp. cuneata]) are found on sandy soils and old stabilized dunes with
102

relatively open vegetation (P. Baye, pers. observ. 1990's). Kellogg’s horkelia is

presumed to be extirpated in San Francisco, Marin, and Alameda Counties (Skinner

and Pavlik 1994), and is rare on sandy soils of the central coast (Monterey, San Luis

Obispo, and Santa Barbara Counties; Hoover 1970, Matthews 1997). One occurrence

in San Mateo County on part of San Bruno Mountain (Daly City, on a remnant sand

slope in a residential area) is presumed to be the last in the San Francisco Bay area

(Skinner and Pavlik 1994). In San Francisco, Kellogg’s horkelia was collected in the

early 20th century (variously as Potentilla lindleyi var. sericea (A. Gray) Howell, or

Potentilla kelloggii Jepson) from dunes of the western part of the City at Point Lobos,

Lake Merced, Sunset Heights, Ocean View (Howell et al. 1958), all known historic

localities of San Francisco lessingia. Additional survey information on this species is

needed in this region.

10. Large-flowered Linanthus, Linanthus grandiflorus (Benth.) E. Greene

Large-flowered linanthus is a small, showy-flowered annual herb in the phlox family

(Polemoniaceae). Large-flowered linanthus has white to pink flowers with short (5 to

6 millimeters, about 0.2 inch) tubes (distinguishing it from common linanthus

[Linanthus parviflorus], another species also found on old dunes), displayed in headlike clusters (Patterson 1993). It is an uncommon species, typically found in sandy

grasslands with low, sparse vegetation. Within the region it is found mainly along

crests of the Santa Cruz Mountains (Thomas 1961) and rarely in Marin County on old

dunes in Point Reyes (Howell 1949), where it still occurs locally in older dune soils,

and associates with the endangered Sonoma spineflower (Chorizanthe valida) (P.

Baye unpublished data 1999) and the rare Marin horkelia (Horkelia marinense). 

Historic San Francisco collections are known from the Presidio (1894) and near Lake

Merced (1891) (Howell et al. 1958, Wood 1996), localities likely to include old dune

or sandy grasslands. No recent collections have been made in San Francisco or San

Bruno Mountain, and it is quite likely now extirpated on the Peninsula. Additional

survey information on this species is needed in this coastal region.

11. Curly-leaved Monardella, Monardella undulata Benth.

Curly-leaved monardella is a strongly scented (pungent pennyroyal-like fragrance)

annual herb in the mint family (Lamiaceae) with sticky-glandular flowerheads and

purple flowers (Figure 9). It is uncommon but locally abundant, usually found along
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the central coast in old dunes, or similar sandy soils in inland and coastal plant

communities. It ranges from Sonoma to Santa Barbara Counties (Munz 1959). The

only coastal population to be recently confirmed north of Monterey Bay in coastal

dunes occurs in coastal dunes at Point Reyes, where it is widespread and locally

abundant in stabilized older dunes, especially in years of high rainfall (P. Baye

unpublished data 1990 to 1999). The only known locality of this species on the San

Francisco Peninsula was in San Francisco, where a single Brandegee collection

(probably late 19th century) was made at Lake Merced dunes near what is now Fort

Funston (Brandegee 1892, Howell et al. 1958). It is now extirpated on the San

Francisco Peninsula, although suitable habitat persists at Fort Funston near the historic

locality.

12. Greene’s Popcornflower, Plagiobothrys reticulatus (Piper) I. M. Johnson var.

rossianorum I. M. Johnson (= Plagiobothrys diffusus E. Greene)

Greene’s popcornflower is a low annual herb in the borage family (Boraginaceae). The

treatment of Plagiobothrys by Messick (1993) in the Jepson Manual interpreted the

endemic San Francisco (Presidio) population of Greene’s popcornflower

(Plagiobothrys diffusus) as a variant within Plagiobothrys reticulatus var.

rossianorum. The taxonomy of this group is problematic. Thomas (1961) placed

Plagiobothrys reticulatus within Allocarya californica, and cited only one locality

north of Santa Cruz; he attributed Plagiobothrys diffusus only to San Francisco. 

Skinner and Pavlik (1994) reported that Greene’s popcornflower is known from only

six localities, in Santa Clara and San Francisco Counties. Whatever the identity of this

ambiguous set of taxa, they appear to be quite rare. Greene’s popcornflower is

extirpated (possibly altogether extinct, depending on taxonomic interpretation) in San

Francisco, where it was reported only from “clayey flats near Mountain Lake” as

recently as the 1930's (Howell et al. 1958, Wood 1996). 

Greene’s popcornflower was reported from southwest Farallon Island from 1978 to

1985, where it occurred in local abundance between residences (apparently extirpated

since then), but only photographs, no voucher specimens, are available to confirm this

occurrence (Point Reyes Bird Observatory file information; M. Coulter, pers. comm.

1997). Southwest Farallon Island was supplied with imported garden soil for

residential gardens during the last century, possibly from the Presidio, which may have

been the source of the population. Though not observed since 1985, it may possibly
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remain as dormant seed in soil there. Recent reports of this species in dune slacks and

other seasonal wetland habitats of Point Reyes await verification. Additional survey

information on this species is needed, particularly Point Reyes and seed bank sampling

from the Farallons.

13. San Francisco Campion, Silene verecunda S. Watson ssp. verecunda

San Francisco campion is a perennial herb in the pink (carnation) family

(Caryophyllaceae). It grows from a branched semi-woody base that bears erect stems

supporting terminal clusters of small but showy white to pink flowers. It occurs only

between San Francisco and Santa Cruz County, and is reported from fewer than 20

localities (Thomas 1961, Skinner and Pavlik 1994). Wilken (1993) in the Jepson

Manual interpreted these plants as a possibly indistinct variant of ssp. platyota, which

is widely distributed. In San Francisco it was historically reported or collected from

Lands End, Sunset District dunes, and Lake Merced (Fort Funston) dunes (Howell et

al. 1958). It is not known to grow on coastal dunes elsewhere in California. Small

native remnant populations currently occur in San Francisco on dunes above Baker

Beach, where the population has been artificially augmented. It also has been

reintroduced to restored dunes near Lobos Creek and upslope from the natural Baker

Beach population. A second population is reported from Mt. Davidson at the

southeastern part of the City (Wood 1996). The type locality of the subspecies is San

Francisco.

San Francisco campion at the Presidio includes the initial populations as follows: one

small patch just south of the ravine within Disturbed Area 3, one small patch east of

Lincoln Boulevard in the lower north Baker Beach terrace, and one small patch shaded

under cypress trees just north of Feral Scrub. Individuals were introduced in 1993

along the sand ladder, the lower slope north of Battery Crosby Road, the Feral Dunes,

and Lobos Dunes. (Craig Middleton, in litt., 2002)

14. Seashore or Coast Starwort, Stellaria littoralis Torrey

Seashore starwort is a coarse colonial perennial herb in the pink (carnation) family

(Caryophyllaceae) (Figure 9). It grows from tough, elastic creeping rhizomes and has

lax, slender, flexible stems that climb over adjacent vegetation. It occurs in moist or

wet sandy coastal soils, mostly in moist dune slacks or seeps in sandstone bluffs and
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ravines. San Francisco is the historic southern limit of the species, which ranges north

to Mendocino County (Smith and Wheeler 1990-1991). It is rare, though locally

abundant, throughout its historic range, and is known from a few populations at Dillon

Beach (Brazil Beach), Point Reyes grassland swales (Howell 1949, P. Baye

unpublished data 1990's), Bodega Head dunes (P. Connors unpublished data 1998,

Best et al. 1996), and Manchester Beach dunes (Mendocino County; Smith and

Wheeler 1990-1991). It is extirpated in San Francisco, where it was historically

recorded from wet sandy soil near Lands End (Howell et al. 1958). These localities

were probably thin dune deposits over seeps in Franciscan bedrock or Colma

formation sands. The nearest populations are at Point Reyes, the type locality and

center of abundance (from Tomales Point to Limantour Estero in dune slacks and wet

grassland swales).

15. Dune Tansy, Tanacetum camphoratum Less.

Dune tansy is a robust, tall, erect, rhizomatous, and strongly scented perennial herb in

the aster family (Asteraceae) (Figure 9). Its lax, coarse, spreading above-ground stems

and below-ground rhizomes form colonies in stable and semi-mobile coastal dunes.

Dune tansy (Tanacetum camphoratum) was distinguished from Douglas’ tansy

(T. douglasii DC.), which ranges from Mendocino County to British Columbia (Munz

1959), by the absence of conspicuous short yellow petal-like ligulate (ray) flowers

(present in populations from Mendocino and north) and the presence of dense whitewoolly and gray-green leaves. The only natural populations of the typical whitewoolly form with highly reduced or absent ray florets today occur in dunes at San

Francisco, Dillon Beach (Marin County) and Manchester Beach (Mendocino County)

(P. Baye unpublished data 1997 to 2001). Dune tansy from San Francisco was

recently introduced at Linda Mar Beach, Pacifica, San Mateo County. Both taxa have

recently been treated as Tanacetum camphoratum without even varietal distinction

(Kyhos and Raven 1982, Smith 1986, McClintock 1993). This taxonomic revision

made the rare white-woolly, “rayless,” geographically discrete southern populations

merely an extension of a single widespread northern coastal species. More detailed

and comprehensive study of variability within and between populations in California

is needed to improve taxonomic understanding of dune tansy. 

Dune tansy was formerly common within the western parts of the San Francisco dune

system, occurring as far east as Buena Vista Park, and west through Golden Gate Park
106

(Brandegee 1892, Howell et al. 1958). It also is evident in historic photographs of

western San Francisco (G. Gaar unpublished photographic archives 1999). Today the

species occurs only on dunes at Baker Beach, sandy slopes near the Cliff House and

Sutro Heights, at all three Sunset Heights dune remnants, and around Fort Funston

dunes. It was recently reintroduced to Crissy Field dunes (P. Baye unpublished data

1997 to 1999).

16. San Francisco Owl’s-clover, Triphysaria floribunda (Benth.) Chuang and

Heckard (= Orthocarpus floribunda Bentham)

San Francisco owl’s-clover is a very rare, showy, annual hemiparasitic member of the

snapdragon family (Scrophulariaceae). Its habitat is coastal grassland, often on

serpentine or sandy soil, in Marin, San Francisco, and San Mateo Counties (Howell

1949, Howell et al. 1958, Thomas 1961, Skinner and Pavlik 1994). In San Francisco,

it had been known until recently only as a small intermittent population near the

Raven’s manzanita parent clone. In some years, no plants appear there, followed by

years of small populations. It was recently rediscovered on an unirrigated portion of a

rough lawn near the Log Cabin, Presidio, on graded serpentine soil (M. Chasse, pers.

comm. 2001), near or on an historic locality. Historically, San Francisco owl’s-clover

was collected in San Francisco above Fort Point, Baker Beach, Presidio (1902), and

Lake Merced (1907) (Howell et al. 1958, Wood 1996). These localities included

coastal dunes, serpentine outcrops, and older sandy soils. Otherwise it is recently

confirmed only from the Point Reyes Peninsula (B. Moritsch, pers. comm. 1999), and

remains precariously close to extinction. The nearest historic populations from San

Bruno Mountain have not been observed since the 1960's (McClintock et al. 1990). 

C. Species of Local and Regional Conservation Significance

The following species are either in decline in a significant portion of their ranges, or

have become extirpated or rare in coastal dune and serpentine outcrop vegetation on

the San Francisco Peninsula. Some local populations may represent significant

ecological or geographic variants of more widespread species. Some taxa are

suspected of being in decline, but lack reliable and recent records of occurrence. 

These plants warrant scrutiny to avoid significant decline in geographic range and

abundance. Conservation of genetic variation among populations of relatively

widespread species is beneficial, and is efficient compared to intervening after a
107

species becomes rare due to artificial factors (Millar and Libby 1991). Like the

species of concern, these selected species of local and regional interest warrant special

attention in surveys and restoration plans that are conducted as recovery actions for

listed species. These selected species in some cases may be considered likely to

become species of concern, at least within the portion of their range covered in this

recovery plan.

1. Pink Sand-verbena (introgressant) Abronia umbellata Lam. ssp. umbellata 

(introgressive with Abronia latifolia) 

Typical pink sand-verbena (Abronia umbellata) is a fleshy annual herbaceous plant in

the four o’clock family (Nyctaginaceae) (Figure 9). It has a prostrate and spreading

growth habit, slightly sinuate green leaves, and abundant umbels of showy pink

flowers. The historic range of typical pink sand-verbena is Sonoma County to Baja

California (Munz 1959, Spellenberg 1993a). The species as a whole is inconsistently

treated in regional floras as either an annual (Munz 1959, Spellenberg 1993a, Skinner

and Pavlik 1994) or perennial (Jepson 1911, Abrams 1944, Munz 1959, Hoover 1970). 

Also, many poorly defined varieties were formerly recognized (Tillett 1967). This

taxonomic ambiguity may be due to confusion between typical populations and

widespread introgressive (natural hybrid backcrosses) populations, often with yellow

sand-verbena (Abronia latifolia) (Tillett 1967). Introgression of yellow sand-verbena

into pink sand-verbena is widespread in California, while introgression of pink sandverbena into yellow sand-verbena is rare (Tillett 1967). 

Introgressive populations of pink sand-verbena have been identified from San

Francisco, San Mateo, Marin, and Sonoma Counties (Tillett 1967, and California

Academy of Sciences herbarium collection annotations by Tillett, 1961). Populations

from San Mateo beaches and low foredunes northward to Marin County are currently

very uncommon and small, and appear to be predominantly perennial introgressive

forms. Typical pink sand-verbena is currently common only in stable dune scrub from

the Monterey Bay area southward (P. Baye, pers. observ. 1995 to 1999). Introgressive

or hybrid populations are commonly facultative perennials and occur on beaches and

foredunes, like yellow sand-verbena, while the nearest populations of typical annual

pink sand-verbena in Monterey Bay usually occur in stabilized older dunes (P. Baye,

pers. observ. 1997 to 1999). Traits that indicate introgression or hybridization with

yellow sand-verbena include fragrant flowers, pale pink to pale salmon or yellowish-
108

pink flowers (sometimes with yellowish eye-spots), perennial growth, and thinner

gray-green leaves. The rare north coast pink sand-verbena (Abronia umbellata ssp.

breviflora [Standley] Munz) of northern California beaches, though treated as a

distinct subspecies, is also considered to be the stabilized product of introgression with

yellow sand-verbena (Tillett 1967).

In San Francisco, “Abronia umbellata” as reported by Brandegee (1892) and Howell

et al. (1958) occurs only at the low foredunes and beach at Crissy Field, in the

Presidio (a site evidently not surveyed by Tillett [1967]). This distinctive population

occurs mixed with yellow sand-verbena, and appears to consist exclusively of

introgressive individuals, and possibly some hybrids. Some individuals here approach

ssp. breviflora (P. Baye unpublished data, 1999). Historically, pink sand-verbena

(probably also introgressant plants) was also reported from a beach in southeastern

San Francisco near Hunters Point (Brandegee 1892). It was also collected from

Ocean Beach near Golden Gate Park by Peter Rubtzoff in 1959 (CAS538598), and

from an unspecified San Francisco locality by Kellogg in 1868-1869 (CAS132847),

annotated by Tillett as a mixed collection of the typical species and introgressants. 

Introgressants have also been collected from Lands End (Tillett 1967).

The introgressant pink sand-verbena of Crissy Field is protected by the Golden Gate

National Recreation Area, and is also propagated and outplanted locally in local dune

restoration (J. Cannon and M. Wadsworth, pers. comm. 1997 to 1999). The natural

population of perennial foredune clones here is highly variable: plants are generally

perennial with wavy-margined gray-green leaves and flowers variously lightly scented

or unscented by day, ranging from pale salmon with yellow highlights and highly

scented to rich magenta with minimal scent. The population is apparently stabilized

and sexually reproducing: it has been present at this locality since the late 19th

century (Brandegee 1892), and seedlings spontaneously occur some years at Crissy

Field (P. Baye, pers. observ. 1993 to 1998). Field-collected seed germinate readily in

the Presidio nursery (M. Wadsworth, pers. comm. 1998). However, nursery

propagation by seed may be unconsciously selective. Nursery-grown transplants

appear to have high frequencies of unscented, deeper pink-flowered genotypes that are

closer to typical pink sand-verbena than the local old intermediate forms. 
109

Most foredune habitat at Ocean Beach is probably too unstable for reintroduction of

pink sand-verbena introgressants, but suitable habitat exists at the restored bluff-top

foredunes at southern end of Fort Funston.

2. Coast Rock-cress, Arabis blepharophylla Hook. & Arn.

Coast rock-cress is a perennial herb in the mustard family (Brassicaceae). It forms

clumps of vegetative rosettes that develop flowering shoots in late winter and early

spring. Selected forms of the species are cultivated as an ornamental perennial for

purple spring-blooming flowers. Coast rock-cress naturally occurs on outcropping

bedrock along or near the coast, or among sparse vegetation or open soil of coastal

bluffs from Santa Cruz to Sonoma Counties. It is uncommon to rare along the bluffs

of the Golden Gate, particularly the Presidio serpentine bluffs (Golden Gate National

Recreation Area, unpublished file information 1995). Local populations occur on San

Bruno Mountain. Coast rock-cress is threatened by invasive species that colonize its

naturally open habitats. Pampas (jubata) grass (Cortaderia jubata) and iceplant are

examples of these aggressive types of invasive species. San Francisco is the probable

type locality.

3. Nuttall’s Milk-vetch, Astragalus nuttallii (Torrey & A. Gray) J. Howell var.

virgatus (A. Gray) Barneby

Nuttall’s milk-vetch is an uncommon to rare taprooted perennial herb in the pea family

(Fabaceae) with low, sprawling or mounding stems (Figure 9). Like other

milkvetches, it produces large, dry, papery-skinned, inflated, buoyant fruits (legumes)

and small, hard seeds. This species typically grows in vegetation of sandy coastal

bluffs, coastal scrub, and coastal dune scrub. The northern limit of the variety virgatus

is Sonoma County (Salt Point, north of Gerstle Cove, where multiple colonies occur in

grassland of a marine terrace near the bluff edge; Best et al. 1996, P. Baye, unpubl.

data 2000). The reported southern limit of this variety is a coastal bluff south of

Pomponio Beach, San Mateo County (CALFLORA 2001). The largest population

occurs at Mori Point, Pacifica, San Mateo County, on ocean bluffs of a derelict quarry

(P. Baye, unpubl. data 2001). Elsewhere in San Mateo County, Nuttall’s milk-vetch

occurs uncommonly in grassland and open sandy soils of San Bruno Mountain

(McClintock et al. 1990), near Sweeney Ridge near Pacifica (M. Mencke, pers. comm.

1998), and on coastal bluffs near Pillar Point, San Mateo County (P. Baye unpublished
110

data 1998 to 2001). It has been reported in Marin County as local and rare in coastal

bluffs at Tomales Point, but has not recently been confirmed there (Howell 1949; P.

Baye, pers. observ. 1990's). Nuttall’s milk-vetch is currently restricted in San

Francisco to Fort Funston dunes and recent transplants to Crissy Field, Presidio (P.

Baye, pers. observ. 2001). Historically it was collected at the Presidio (1900, 1894),

Point Lobos (1912), Lone Mountain, and Lake Merced (1892) (Howell et al. 1958,

Wood 1996). San Francisco is the type locality of Franciscan milkvetch (Astragalus

franciscanus [Sheldon]), which has been placed in synonomy with Astragalus nuttallii

var. virgatus. The similar variety nuttallii (distinguished by its grayish hairy leaves

and looser inflorescence; Munz 1959, Spellenberg 1993b) ranges from Monterey Bay

south to Point Conception (Munz 1959).

4. California Saltbush, Atriplex californica Moquin

California saltbush is a low perennial herb in the goosefoot family (Chenopodiaceae). 

It grows in clumps or mats branching from a thick, deep taproot, and has small lanceshaped gray-green to greenish white-mealy leaves. Its habitats include coastal dunes,

coastal bluffs, and coastal salt marsh edges. It is uncommon north of Monterey Bay,

ranging north to northwest Sonoma County (Taylor and Wilken 1993, Best et al.

1996), where a small, usually vegetative colony grows on sandstone outcrops in

coastal cliffs near Gerstle Cove, Salt Point. In west Marin County, it generally occurs

along the upper edges of sandy salt marshes, and on coastal sandstone bluffs. From

Monterey Bay south, it occurs in stable dunes and sandy bluffs in areas of

predominantly low vegetation. Historically it occurred on bluffs and dune flats in

western San Francisco (Howell et al. 1958). Locally it is currently restricted to small

remnant populations at Fort Point (on erosional serpentine bluffs, an atypical habitat)

and larger populations near Point Lobos (on Colma Formation sands and Franciscan

sandstone). It is extirpated in San Francisco dune remnants and salt marsh edges, and

is not reported from dunes north of Monterey Bay. Nearest recently confirmed

populations occur on sandy bluffs in San Mateo County (Rockaway Head and Mori

Point, Pacifica) and along salt marsh edges at Point Reyes (P. Baye unpublished data

1997 to 2001). A natural population occurs just east of the Coast Guard station in the

foredune area, which has now expanded through Crissy Field restoration efforts 

(Craig Middleton, in litt., 2002).
111

5. Indian Paintbrush, Castilleja affinis Hook. and Arn. ssp. affinis (coastal forms)

The taxonomy and identification of variable coastal Indian paintbrush populations

(Castilleja spp.) is complex. The Indian paintbrush of the serpentine bluffs and dunes

of San Francisco was treated as Monterey Indian paintbrush (Castilleja latifolia) in the

Flora of San Francisco (Howell et al. 1958). Typical C. latifolia has been narrowly

interpreted as a species restricted to the vicinity of the Monterey Bay/Santa Cruz coast

(Pennell 1951, Munz 1959, Chuang and Heckard 1993), but it also has been broadly

interpreted as a variable species with many infraspecific taxa distributed over a wider

geographic range (Jepson 1925, Howell 1949, Howell et al. 1958, Thomas 1961),

including many taxa now placed in synonymy with coast Indian paintbrush (C. affinis)

or Wight’s Indian paintbrush (C. wightii) (Chuang and Heckard 1993). The taxonomy

of all the perennial Castillejas of the immediate central and northern California coast

(C. latifolia and its former varieties, C. wightii, C. mendocinensis, C. affinis ssp.

litoralis and ssp. affinis [including C. inflata Pennell]) has been variously interpreted

throughout the 20th century (Jepson 1925, Howell 1949, Pennell 1951, Munz 1959,

Chuang and Heckard 1993). The notorious taxonomic difficulties in the genus are due

to high variability within species, hybridization, and polyploidy (Chuang and Heckard

1993).

Following the revised key to species of the realigned Castilleja by Chuang and

Heckard (in Hickman 1993), plants from coastal bluffs and dunes in San Francisco,

and herbarium sheets of most “Castilleja latifolia” from San Francisco at the

California Academy of Sciences would be referred to coast Indian paintbrush

(C. affinis ssp. affinis), but possibly intergrading toward Monterey Indian paintbrush

(C. latifolia) in some significant traits (P. Baye unpublished data 1999) (Figure 9). 

Intermediate plants referable to C. affinis, but closely approaching typical Monterey

Indian-paintbrush are prevalent in dunes at Pescadero, San Mateo County (P. Baye

unpublished data 1998 to 2001). The San Francisco (and Marin) coastal populations

of coast Indian paintbrush are atypical for this species in significant vegetative

morphological (mostly foliar) traits. Presidio serpentine bluff and Baker Beach dune

populations appear morphologically indistinct. Presidio serpentine bluff and dune

populations of coast Indian paintbrush have firm, thick membranous leaves (not fleshy

as typical Monterey Indian paintbrush) that are mostly lanceolate (occasionally lanceovate to oblong, much broader than typical coast Indian paintbrush), up to 7

centimeters (2.8 inches) long (cauline leaves typically 3 to 6 centimeters long [1.2 to
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2.4 inches]), acute, more than 3 times longer than wide (often more than 5 times

longer than wide), often involute or folded (abaxial surface outward), with relatively

narrow to linear, acute lateral or terminal lobes up to one-third the length of the leaf

(P. Baye unpublished data). 

The San Francisco serpentine bluff and dune populations of coast Indian paintbrush

are significant for their size, atypical traits, and wide ecological tolerance for extremes

of exposure to salt spray and harsh, contrasting soil conditions. They apparently

displace the common Wight’s Indian paintbrush, the dominant Indian paintbrush of the

northern San Mateo coastal bluffs and headlands. The occurrence of coast Indian

paintbrush on serpentine soil here is significant: no Monterey Indian paintbrush

collections from serpentine are reported (CALFLORA 1999), but another subspecies

of C. affinis, the Tiburon Indian paintbrush, federally listed as endangered, (C. affinis

ssp. neglecta; U.S. Fish and Wildlife Service 1998a) is restricted to serpentine soils in

Tiburon, less than 8 kilometers (5 miles) from the Golden Gate.

Howell (1949) also treated the similar coastal dune Indian paintbrushes of Point

Reyes, formerly classified as Castilleja inflata Pennell, as variants of C. latifolia. 

Although much of the “Castilleja inflata” from spray-exposed Point Reyes localities

more closely approaches Monterey C. latifolia in critical vegetative traits (oblong to

ovate, blunt, succulent leaves; P. Baye unpublished data), Chuang and Heckard (1993)

placed C. inflata in synonymy with C. affinis ssp. affinis. C. affinis ssp. affinis

specimens similar to those of San Francisco also occur in coastal scrub at Point Reyes. 

Geographic and environmental variation in vegetative characters in the coastal

C. affinis - latifolia - wightii - mendocinensis complex may require more intensive

sampling of populations for adequate taxonomic resolution.

Coast Indian paintbrush (C. affinis ssp. affinis) is a hemiparasitic perennial herb with a

woody base, and a member of the snapdragon family (Scrophulariaceae). Coast Indian

paintbrush grows in clumps (sometimes decumbent to ascending) about 0.3 meter (1

foot) tall in open coastal scrub vegetation on dunes and bluffs in San Francisco. It

typically has abundant short axillary shoots on the coast from San Mateo County to

Marin County. In late summer or fall after seeds have matured, it dies back to a

woody base, and regenerates vegetatively during late winter when soil moisture,

temperatures, and daylength increase. Seedlings often occur shaded under parent

plants or leaf litter in years of ample rainfall. Nearly glandless (below the
113

inflorescence, unlike glandular Wight’s Indian-paintbrush), shaggy-hairy or bristly

stems bear spikes of flowers beginning around March to April, and continue to appear

intermittently through fall or winter. Conspicuous bracts (sheath-like leafy structures

appearing below flowers) range in color from dull orange-red to brilliant scarlet in San

Francisco; elsewhere they may occasionally include orange-yellow to salmon-yellow

forms as well. Hummingbirds are presumably the principal pollinators. 

Coast Indian paintbrush in San Francisco was reported historically as Castilleja

latifolia from dunes and bluffs above Baker Beach, around Lake Merced (now Fort

Funston area), Presidio (serpentine) bluffs from Fort Point to Lobos Creek, and from

Sunset District dunes (Brandegee 1892, Howell et al. 1958). Modern dune

populations are limited to Baker Beach (1 colony of fewer than 50 plants), Hawk Hill

at Sunset Heights (approximately 4 plants in 1999), and restored and relict dunes of

Fort Funston (variable populations, up to several hundred). The Presidio serpentine

bluff population is relatively large (hundreds of plants some years). Population size

varies significantly among years, but does not fluctuate as much as annual Castilleja

species (P. Baye, pers. observ. 1990 to 1999). Coast Indian paintbrush has been

propagated and planted at Fort Funston and Presidio dunes by the Golden Gate

National Recreation Area (M. Petrelli and M. Wadsworth, pers. comm. 1997, 1998).

According to Pennell (1951) the type locality of coast Indian paintbrush is presumed

to be San Francisco, and the type locality of Monterey Indian paintbrush is Monterey,

but a San Francisco type locality of Monterey Indian paintbrush also has been

attributed to Hooker and Arnott (Howell et al. 1958). 

6. Banded (Broadleaf) Purple Owl’s-clover, Castilleja exserta (A.A. Heller) Chuang

and Heckard ssp. latifolia (S. Watson) Chuang and Heckard

Banded purple owl’s-clover is a showy annual hemiparasitic herb in the snapdragon

family (Scrophulariaceae) with spikes of purple, white, and yellow flowers (Figure 9). 

The subspecies latifolia (literally, “broadleaf”) is restricted to sandy soils of coastal

bluffs, coastal grasslands, and dunes of the north and central coast of California

(Howell 1949, Thomas 1961, Best et al. 1996, Matthews 1997). It is now uncommon,

but locally abundant over its range. In San Francisco, it is apparently either rare,

intermittent (emerging only some years), or extirpated in coastal bluffs and dunes. 

Historically it was locally common at Lake Merced (1891, and as recently as the
114

1950's), and Ocean View (Howell et al. 1958, Wood 1996). It is reported as frequent

in grasslands of San Bruno Mountain (McClintock et al. 1990). The nearest

population on the immediate coast, which may be related to C. exserta ssp. latifolia

(possibly related to C. ambigua) is at Rockaway Head, Pacifica (headland south of

Rockaway Beach), where an anomalous form grows near the edges of a thin old dune

soil (P. Baye unpublished data 1999). Banded purple owl’s-clover also occurs in

coastal dune grassland (older dune soils) at Point Reyes (south end of the outer dune

system; Howell 1949, P. Baye unpublished data 1997 to 1999) and near Marina,

Monterey Bay (Matthews 1997; P. Baye, pers. observ. 1997 to 1999). The species is a

host plant for larvae of the threatened bay checkerspot butterfly (Euphydryas editha

bayensis; U.S. Fish and Wildlife Service 1998b).

7. California Goosefoot, Chenopodium californicum (S. Watson) S. Watson

California goosefoot is a robust, coarse-leaved taprooted perennial herb in the

goosefoot family (Chenopodiaceae). It occurs in a wide range of plant communities in

relatively dry, open conditions. California goosefoot is infrequent to uncommon in

central coastal scrub, but is widespread in California west of the Sierra (Munz 1959). 

It was historically uncommon or rare on the San Francisco Peninsula and San Bruno

Mountain (Howell et al. 1958, McClintock et al. 1990). California goosefoot

populations occur in dune areas in the Presidio just north of Battery Crosby road, in

dune scrub at Rob Hill (M. Chasse, pers. comm. 1998), above Baker Beach (this

population has been enhanced by supplemental plantings), and on the south side of the

Presidio Hills restoration site (Brian O’Neill, National Park Service, in litt. 2002). 

California goosefoot populations include a small patch that has been intermittently

observed on the northern side of the trail traversing south of the Presidio Hills site

(Craig Middleton, in litt., 2002). A small planted population has been established in

the dune remnant “Quail Commons” site, southeast of the intersection of Battery

Caulfield Rd. and Washington Blvd (Brian O’Neill, National Park Service, in litt.

2002). Historic records in San Francisco are limited to the Presidio, and a sandy

hollow east of Lake Merced (Howell et al. 1958). The nearest known coastal dune

populations are from San Mateo County (Half Moon Bay) and Point Reyes (P. Baye

unpublished data 1998 to 1999). It is relatively more frequent in stable old dunes of

Monterey Bay.
115

8. Davy’s Clarkia, Clarkia davyi (Jepson) Harlan Lewis and M. Lewis

Davy’s clarkia is an annual member of the evening-primrose family (Onagraceae). It

has showy lavender-rose flowers, and a growth habit ranging from prostrate to

ascending. Historically Davy’s clarkia was known from dunes of the Sunset district,

Lake Merced, and near Ocean View (Howell et al. 1958). It ranges from Del Norte

County to San Mateo County along the coastal bluffs and sands (Munz 1959). It is

more frequent in the northern part of its range. Its southern limit along the coast was

reported by Howell et al. (1958) to be San Francisco, although it is reported as

occasional on San Bruno Mountain (McClintock et al. 1990), and a disjunct

population is reported from Santa Rosa Island (Lewis 1993).

9. California Croton, Croton californicus Muell. Arg.

California croton is a prostrate gray-green subshrub or shrub in the spurge family

(Euphorbiaceae). It occurs on sandy soils (dunes, inland sand deposits, alluvial fans)

from San Francisco south to Baja California, and inland to Arizona (Webster 1993). It

is common in coastal dune scrub along the southern central coast (P. Baye unpublished

data 1995 to 1999). North of the Monterey Bay region it is rare on the coast, collected

or reported historically only from San Francisco (Thomas 1961). The northern coastal

limit is in the remnant dunes of Presidio. It also occurs in remnant dune scrub at

Sunset Heights (Hawk Hill, 14th Avenue near Riviera Street). The nearest coastal

populations are in Monterey County; the nearest interior populations are at the

Antioch Dunes, Contra Costa County (Munz 1959). San Francisco is the type locality.

10. Franciscan Leafy-daisy, Erigeron foliosus Nutt. var. franciscensis G. Nesom

Franciscan leafy-daisy is a perennial herb in the aster family (Asteraceae) that grows

from a low woody base, with daisy-like flowers and narrow linear leaves. It occurs in

sandy grasslands or openings in coastal scrub of stable dunes, bluffs, or woodlands. 

Regionally, the species was uncommonly collected in San Mateo and Marin Counties

(Howell 1949, Thomas 1961), but these floras assigned the local taxa to Erigeron

foliosus var. hartwegii. Erigeron foliosus var. hartwegii, as currently interpreted,

occurs only in the north and central Sierra, while all Erigeron foliosus in the San

Francisco Bay area is referred to variety franciscensis (Nesom 1993). In San

Francisco, leafy-daisy occurred historically at Point Lobos and Lake Merced (Howell
116

et al. 1958), both likely coastal dune scrub habitats. Its current status in San Francisco

is not known, but it has not been recently reported, and may be locally extirpated.

11. Yarrow-leaf Gilia, Gilia millefoliata Fischer and C. Meyer 

Yarrow-leaf gilia (Gilia millefoliata) is an annual herb in the phlox family

(Polemoniaceae) with a skunk-like scent, finely dissected leaves, and sparse clusters

(not dense heads) of purplish flowers. It occurs generally on stable coastal dunes,

becoming relatively frequent on north coast dunes from Mendocino to southern

Oregon (Smith and Wheeler 1990-1991, Day 1993). The taxonomic affinity of the

Gilia millefoliata of San Francisco is questionable. All San Francisco collections were

made in rocky grassland habitats remote from coastal dunes, often in serpentine

(Howell et al. 1958). Thomas (1961) and Howell et al. (1958), placed the San

Francisco collections within the more widespread Gilia clivorum (Jepson) V. Grant, a

species common in grassy vegetation of rocky soils, but these specimens were

interpreted as Gilia millefoliata in San Francisco by Day in the Jepson manual

(Hickman 1993), despite the anomalous soil type and habitat for Gilia millefoliata, but

typical habitat for Gilia clivorum. Day’s key distinguishes these two taxa based on

continuous variation in calyx length; Munz’s [1959] key distinguishes them using

qualitative characters, but excludes Gilia millefoliata from south of San Francisco, and

attributes it solely to dunes. The San Francisco populations are apparently extirpated. 

Yarrow-leaf gilia likely may have occurred in San Francisco dunes as well, but Howell

(1949) stated that it occurred no farther south than Point Reyes. Classification of the

extirpated San Francisco plants should be verified from herbarium specimens, since

the historic rocky habitats of San Francisco collections are most congruent with the

common and variable Gilia clivorum of the coast ranges.

12. San Francisco (Chamisso’s) Dune Gilia, Gilia capitata Sims ssp. chamissonis (E.

Greene) V. Grant

San Francisco dune gilia (colloquially “dune gilia”) is another skunky-scented annual

herb in the phlox family (Polemoniaceae) (Day 1993) (Figure 9). It is restricted

mostly to vegetation gaps in low-growing coastal dune scrub and stable dune

grassland. Dune gilia has showy clusters of lavender-blue to deep violet flowers borne

on long stalks, and has finely dissected linear leaves that are sometimes fleshy. It

occurs infrequently but in local abundance in sparse coastal dune scrub and grassland
117

vegetation from Sonoma County (Best et al. 1996) to San Mateo County (including

Hillside Park, Daly City, on inland sands with San Francisco lessingia) (Howell 1949,

Howell et al. 1958, Thomas 1961, McClintock et al. 1990). In San Francisco it

occurs in most large dune remnants, and also on sandy soils of Yerba Buena Island

(M. Wood, pers. comm. 1999). San Francisco forms appear to be distinctly taller, and

have less intense purple pigmentation than forms from the largest of the subspecies’

populations at Point Reyes dunes (P. Baye, pers. observ.). The type locality of Gilia

achilleifolia ssp. chamissonis (Greene) Brand and other synonyms of this subspecies

(but not of the species Gilia capitata, in which this taxon is now placed) is San

Francisco.

13. Wedge-leaf Horkelia, Horkelia cuneata Lindley ssp. cuneata

 Like the related Kellogg’s horkelia (H. cuneata var. sericea), wedge-leaf horkelia is

an herbaceous perennial member of the rose family (Rosaceae). In dune systems,

wedge-leaf horkelia is generally restricted to older dune soils (P. Baye, pers. observ.). 

It is more widespread and common in sandy coastal soils than Kellogg’s horkelia

(Thomas 1961, Ertter 1993), but it has apparently become extirpated in San Francisco

dunes, and is rare on coastal dunes north of Monterey Bay (P. Baye, pers. observ.

1990's). Wedge-leaf horkelia typically grows on old dunes and sandy soils in coastal

grassland or scrub vegetation along the central and southern California coast (Munz

1959).

14. Common Linanthus, Linanthus parviflorus (Bentham) E. Greene

Common linanthus is a relatively more common showy annual herb in the phlox

family (Polemoniaceae), compared with its relative, Linanthus grandiflorus (Patterson

1993) (Figure 9). It is widespread in grasslands, sandy soils, and partly shaded rocky

slopes in the San Francisco Bay area and the north coast ranges. The nearest coastal

dune populations are known from Point Reyes, where it occurs in old dune soils with

the endangered Sonoma spineflower (Chorizanthe valida) and the rare Marin horkelia

(Horkelia marinensis) (P. Baye unpublished data 1999). It was formerly collected in

colonies at the Presidio and Lake Merced (probable dune localities) from the late 19th

century to the middle 20th century (Howell et al. 1958), and was widespread in stable

dunes of the western part of the City in the late 19th century (Brandegee 1892). It has

been locally extirpated in remnant dunes of San Francisco. It was reported in the San
118

Francisco flora (Howell et al. 1958) as Linanthus androsaceus (Benth.) E. Greene var.

croceus (Milliken) H. Mason, a taxon now placed in synonymy with Linanthus

androsaceus (Patterson 1993) as common on coastal bluffs and grassy slopes of the

northern San Mateo coast (Thomas 1961).

15. Skunkweed, Navarretia squarrosa (Eschscholtz) Hook. and Arn.

Skunkweed is a common and widespread annual member of the phlox family

(Polemoniaceae), easily recognized by its resinous foliage and spiny flower-heads

with strong skunky glandular scent. Skunkweed is generally common in sandy

alluvium, roadsides, dried winter pools, and general disturbed annual habitats. It was

formerly found in dunes and sandy soils of the Sunset, Lake Merced, and the Presidio

(Howell et al. 1958, Wood 1996), but locally has become rare in San Francisco’s

remnant dunes and sandy soils of the Presidio (one site off Battery Caulfied Road). 

The Presidio of San Francisco was the type locality (Howell et al. 1958). 

16. California Broomrape, Orobanche californica Cham. and Schldl. ssp. californica

 California broomrape is a parasitic, leafless, nongreen plant in the broomrape family

(Orobanchaceae) is generally uncommon (Heckard 1993). Clustered broomrape,

Orobanche fasciculata Nutt., a widespread species, is local and uncommon as well;

neither has been reported or collected from San Francisco in recent years. California

broomrape appears occasionally in dunes from Goat Rock (Russian River mouth,

Sonoma County) to Morro Bay dunes (San Luis Obispo County), and it also occurs in

coastal bluff grassland (P. Baye unpublished data 1990's). California broomrape was

collected from San Francisco sand dunes (locality unspecified) early in the 20th

century (Howell et al. 1958). Clustered broomrape has been collected from serpentine

vegetation on the Presidio and Laurel Hill Cemetery, historic localities of Raven’s

manzanita. Little is known about the propagation or reintroduction of these obligate

parasitic plants. San Francisco is the type locality of California broomrape.

17. Coast Rein-orchid or Coast Piperia, Piperia elegans (Lindley) Rydb. 

Coast rein-orchid is a terrestrial orchid that grows leaves in spring, dies back in early

summer, and produces a nongreen spike of white flowers in late summer and fall

(Figure 9). It is uncommon and local on sandy coastal bluff grassland and scrub in the
119

Presidio, and under blue gum groves in remnant dunes near sea level behind Baker

Beach (Golden Gate National Recreation Area unpublished data 1998). It also is

historically uncommon in coastal scrub of the San Mateo coast (Thomas 1961) and

rare on the Marin coast (Howell 1949). It was formerly common in the dunedominated western parts of San Francisco earlier this century (Howell et al. 1958). 

San Francisco was the type locality of the synonym Habenaria greenei Jepson. 
120

IV. RECOVERY

A. Objectives and Criteria

The overall objectives of this recovery plan are to ameliorate or eliminate the threats

in order to improve the population and habitat status of the species sufficiently to

warrant delisting of San Francisco lessingia and reclassification of Raven’s manzanita

to threatened status, and to prevent the species of concern from becoming threatened

or endangered. (Recovery of Raven’s manzanita sufficient to warrant delisting is not

projected for the foreseeable future.) Instrumental to these objectives are the

protection and stabilization of existing populations, and the expansion of restored

natural communities with augmented and newly established populations of listed

species and species of concern.

This recovery plan’s recommendations are based on current understanding of the

biology of the species. The recovery criteria are based on the biology of the species,

the threats to the species, and the strategy that is recommended to counter the threats. 

Information about the recovery strategies used to develop the recovery criteria is

provided in the species account sections in Chapter II. 

Recovered species are expected to have been restored to a point where their long-term

survival in nature is ensured. Criteria used to evaluate when listed species are

recovered should include number and distribution of populations, population sizes, and

probabilities of persistence over specific time periods (Mace and Lande 1991, Tear et

al. 1993, Schemske et al. 1994, Carroll et al. 1996).

More and better data increase the reliability of population forecasting and assessment

of recovery potential and methods (Scott et al. 1995). Development of recovery

criteria for endangered species is often limited by the availability of resources with

which to obtain relevant scientific information (National Research Council 1995). 

Because new data may change our appraisal of what constitute appropriate recovery

criteria, the criteria recommended in this plan are preliminary, and warrant

reevaluation when additional data become available.
121

Future modifications of recovery actions will be informed by research results and

management experience that follow from implementation of recovery actions. This

adaptive management approach will build on successful demonstration of the

functions of the dune scrub ecosystem, as measured by satisfactory maintenance of

populations of San Francisco lessingia and co-occurring species of concern following

their re-establishment across the recovery unit. 

This adaptive management approach will be applied to the areas where the plan has

provided for interim management of portions of the Presidio Recovery Unit to allow

for avoiding impacts to other resource values, as well as areas where we recommend

establishing long term management for San Francisco lessingia immediately.

We recognize the implementation of recovery plans is always affected by

circumstances and new information. Further, this recovery plan focuses on biological

analysis— review of other aspects of implementing the recovery actions in the plan

will be done as required by such laws as the National Environmental Policy Act and

the National Historic Preservation Act, as the Presidio is listed on the National

Register of Historic Places, and parts are designated as National Historic Landmark

Districts. If requested funding is not approved, and other partnerships cannot be

formed, the schedule will have to be adjusted to the means available.

1. Recovery Criteria for San Francisco Lessingia

The strategy for recovery of San Francisco lessingia relies on the establishment of a

large enough area to minimize the edge effects of the adjacent vegetation, including

shading; too much blocking of wind; and spreading of invasive weed species via

seeds, rhizomes, and layering. Core populations at the larger habitat areas are to be

supplemented by smaller, satellite populations. The smaller area available for

management of the satellite populations means that edge effects will be greater;

elimination of invasive species will likely not be possible, even in the center; and they

may blink out from time to time, necessitating re-establishment from seed stores. 

While important to recovery, the satellite populations cannot be relied on for more

than a supporting role in achieving recovery in the wild. Any significant reductions in

the size of the larger habitat areas proposed would result in an unacceptable reduction

in the likelihood of recovery in the wild.
122

The strategy for recovery of San Francisco lessingia, specifically, has several aspects

that involve an adaptive management approach. Adaptive management is the process

of evaluating and adjusting management as needed. With adaptive management,

objectives are developed to describe the desired condition, management is designed to

meet the objectives, the response of the resource is monitored to determine if the

objective is being met, and management is changed if the objectives are not met

(Elzinga et al. 1998). Results of new biological research are considered in adaptive

management. 

The recommendations for invasive cover are our best estimate of the levels needed to

minimize the need for large amounts of hand-weeding. The weed management

recommendations are based on the principles of weed management that have been

shown to be the most efficient (Dewey 1995). The weed management principles call

for prompt identification of colonies of an invasive species that can be treated for

eradication rather than treating entire areas for suppression. Persistence of the

invasive in very low numbers is allowed over large areas, but specific colonies or

areas of invasion are targeted for treatment. These principles are expected to be

applied to newly restored areas that begin with low or no invasive species cover, and

to areas where invasive species have been essentially eradicated. For the core

populations on larger areas of restored habitat, the intent is that most of the invasive

cover be kept from moving inside the outer margins of the restored dune scrub habitat. 

This will change the weed control need from suppression over the whole site to

suppression at the edges and removal of new colonists in the core area before they can

spread and require whole site treatment. Experience with new methods may show that

somewhat higher or lower levels of invasive species as proportions of total cover and

plant cover may be the most efficient means to maintain dune scrub habitat. In this

case the new information would provide the basis for future management.

We, the U.S. Fish and Wildlife Service, recommend that restoration of dune scrub, in

areas of the historic forest within the San Francisco lessingia Presidio Recovery Unit

where many of the existing trees are dying. This restoration should be initiated,

following completion of all environmental review processes, in accordance with

management plans developed and approved by the Golden Gate National Recreation

Area and the Presidio Trust. However, we recommend any replacement of dead trees

should be conducted with dune scrub habitat trees and other perennial dune scrub

species, such as, Coast live oak (Quercus agrifolia), narrow-leaved willow (Salix
123

exigua), and arroyo willow (Salix lasiolepis). (See Appendix I. for a complete list of

native dune scrub plant species.) Each species should be planted at densities

equivalent to the naturally occurring densities found in dune scrub habitat. This

replacement technique should minimize interference with the dune scrub restoration

process. It will also reduce the aesthetic impact of removal of dead trees by giving the

replacements a chance to grow before the remaining trees are removed in later phases

of habitat restoration over the next 30 years. This document describes adaptive

management processes to determine appropriate management of historic forest within

several portions of the Presidio Recovery Unit.

Rob Hill Forest Buffer and Public Health Services Hospital Buffer are two areas

containing relictual dune substrate, adjacent to the recovery unit, with lower suitability

for management as habitat to support San Francisco lessingia. (Please see Figure 11.) 

These areas are on the backslope or leeward side of a dune, rather than the foreslope or

windward side. Management using trees and shrubs along the upper slopes on the

backslope may benefit the dune scrub habitat area by acting as a buffer against

invasion by weedy species, without diminishing the effective functional area of the

core habitat for San Francisco lessingia. Native, low-growing trees and shrubs,

arranged to avoid blocking winds over the top of the dune and the dropping of litter on

the foredune side, with the use of species that do not need watering and fertilization

would inhibit weeds next to the dune scrub habitat. Watering during initial

establishment of new plants is acceptable, but long term watering that increases weed

establishment in the buffer areas or the dune scrub habitat is to be avoided. Regular

monitoring and removal of weeds in these management zones will further inhibit

movement of nonnative species into the areas of dune scrub habitat. This would

support the overall goal of establishing an area of functional dune scrub habitat that

would be largely self-sustaining and support adequate population levels of the species

in the areas that are made available by recreational disturbance and patches of blowout in the matrix of dune scrub vegetation. In this way, the community is selfsustaining and has replacement in perpetuity of San Francisco lessingia individuals in

the population through natural reproduction. 

Disturbance is important to maintaining the dune scrub community. Recreational

disturbance, wind, grazing by goats, and by other methods may all be useful to prevent

a degree of canopy closure that would impair recruitment of San Francisco lessingia

and similar dune scrub species. Due to the importance of the Presidio Recovery Unit

to recovery in the wild, it will be necessary to monitor, and maintain, just enough
124

disturbance to keep it suitable for the shorter-lived plant species, without destabilizing

the dune structure. Results from monitoring will be used to adaptively manage the

habitat to meet the recovery criteria.

Some portions of the recovery unit proposed for restoration will take extra steps, and

time, to re-establish the dune scrub ecosystem due to the increased content of organic

matter in the soil from the non-native trees. We believe that the organic matter will be

reduced to suitable levels by oxidation and leaching if the landscape species (i.e.,

trees) are removed. This process could be accelerated by various management

treatments. The use of dune scrub perennial plants at higher than normal densities

may be needed to bridge the period of soil rehabilitation to block weed entry. 

Feasibility of this area for restoration, and the methods to be used, will be determined

in coordination with the Presidio Trust and Golden Gate National Recreation Area

following detailed analysis of site-specific information during implementation.

The Presidio Trust has suggested that the area of West Washington Housing may be

suitable to add to the Presidio Recovery Unit. The area is adjacent to the Rob Hill and

Wherry Housing sites. We plan to evaluate the site during implementation of this

recovery plan to determine whether it will contribute to the area of continuous

functional habitat that will reduce edge effects on the San Francisco lessingia

population.

Where infrastructure exists (e.g., Building 1750, coastal fortifications, and associated

earthworks), it may be appropriate to maintain a buffer of (preferably low-stature)

landscape vegetation of 15 to 30 meters (50 to 100 feet) around the structures in order

to minimize the likelihood for management conflicts (e.g., building maintenance that

would harm San Francisco lessingia plants if they grew directly adjacent to the

building) due to colonization by San Francisco lessingia. San Francisco lessingia

currently exists in the vicinity of several buildings and structures (e.g., Public Health

Services Hospital, historic batteries, and associated earthworks of the batteries) at the

Presidio, and future management and building maintenance needs for these areas will

likely be dealt with via a programmatic formal consultation under section 7 of the

Endangered Species Act. Consequently, we recommend that the Presidio Trust and

the National Park Service develop a programmatic strategy to manage the areas around

structures, both where San Francisco lessingia now exists and where landscape buffers

are to be maintained. The strategy would describe the areas that require landscape
125

buffers and the management to be done in those areas to avoid adverse effects to the

adjoining dune scrub areas. Please note that the terms “buildings”, “infrastructure”,

and “structures” may include the earthworks around the gun batteries, and other

similar artificial topography that was “built” to fulfill the organic functions of the

military installations at the Presidio, and any prehistoric features as well (such as

hunting blinds, middens, or any mounds, holes, or trenches associated with campsites,

burial grounds or other human use sites).

Reducing the density of the historic forest stands, while leaving them essentially the

same size, may be adequate to allow for the wind disturbance needed to manage

ecologically functional dune scrub habitat. However, the importance of the area of

continuous habitat to be managed as dune scrub must be considered during

management decisions. This is due to the need to minimize edge effects, particularly

by encroachment of non-dune scrub plants into the managed area. Reduced density of

trees along Lincoln Boulevard, together with restoration of a sufficient area to

minimize the need for weed management by extensive hand weeding may be adequate

to support a population level (through fluctuations due to weather and plant

succession) to meet the recovery criteria in the recovery plan. This can only be

determined by establishing a much larger area of dune scrub habitat than currently

exists, followed by monitoring of the San Francisco lessingia population, the bare

ground component, and the vegetation status in the managed area. Restoration is

recommended on the entire recovery unit as configured in the recovery plan, until we

have data showing that ecological functions that maintain dune scrub habitat can

persist over the long term, with retention of the historic forest in the Building 1750

and Bowley Street Interim Management Area. Completion of the restoration in the

area of Lobos Dunes, Wherry Housing, perhaps West Washington Housing, Rob Hill,

and Upper Baker Beach is necessary before we can determine how successful

restoration of function will be, and before deciding that additional restoration of dune

scrub will not be necessary in the Building 1750 and Bowley Street Interim

Management Area and in the Special Management Zone.

The Building 1750 and Bowley Street Interim Management Area will continue, for the

time being, to consist of forest as identified in the Presidio Trust’s and National Park

Service’s Presidio Vegetation Management Plan. Forest management prescriptions

include strict controls on invasive weeds and replacing the current stand at densities

that will allow for healthier trees. However, forest management practices will also be
126

altered to promote wind movement. The area behind Building 1750 and the forested

strip between Lincoln Boulevard and Bowley Street will be converted to lower-stature

trees to allow for more wind passage. Reforestation in these areas will be configured

in thin strips to limit the potential effects of edge and of reduced wind on the valley’s

northern and eastern slopes. Reforestation will be performed with increased spacing

to allow larger openings for wind passage. The density of lower limbs will be reduced

to promote more wind movement. If the monitoring of the Presidio Recovery Unit

shows adequate function in terms of San Francisco lessingia population dynamics and

ecosystem function, with sustainable management methods in the dune scrub (rather

than the intensive methods currently needed, such as extensive hand-weeding), the

long range recommendation in the plan for restoration of dune scrub within the

Building 1750 and Bowley Street Interim Management Area, and away from the

structures could be reconsidered.

Recovery in the Presidio Recovery Unit requires that the San Francisco lessingia

become established in a continuous population, linking the Lobos Creek area with

other existing populations. At the same time, historic preservation considerations for

the San Francisco Presidio, as a National Monument, make it important to consider

ways to minimize the reduction of the extent of the historic forest in the Lobos Creek

area, particularly in the area of the trail running east over the shoulder of the ridge. 

Therefore, an action has been included in the stepdown narrative and implementation

schedule to develop a strategy to address both of these needs. The goal is to

determine, by monitoring, the degree to which the character of the historic forest can

be maintained while allowing recovery and delisting of San Francisco lessingia.

Recovery criteria for San Francisco lessingia apply to existing populations (interim

criteria), to future expanded populations in restored habitats within recovery units, and

to the ecological processes and conditions of the plant communities in which they

occur. San Francisco lessingia may be considered for reclassification to threatened

status when all interim recovery criteria are fully achieved, and expansion of the

Lobos Dunes unit to at least Battery Caulfield Road and upper Baker Beach has been

achieved. The species may be considered for delisting when all long-term recovery

criteria have been met, which could occur no sooner than 20 years after initiation of

implementation.
127

a. Interim Recovery Criteria

Interim recovery criteria apply to existing populations in relict and restored habitats. 

They are aimed at ensuring that these populations do not undergo significant long-term

declines in size, number of subpopulations, areal extent, or population variability

while long-term recovery measures are in preparation. These criteria are primarily

aimed at specific areas occupied by San Francisco lessingia, and the dynamic quality

of the vegetation in these areas. The dynamic quality of vegetation partly determines

the suitability of an area for the ecological regeneration of San Francisco lessingia. 

Progressive trends toward dense, closed woody or perennial vegetation with few open

gaps dominated by annuals would indicate a loss of suitable local conditions for San

Francisco lessingia (see discussion in Chapter II under Ecology and Reproduction). In

contrast, a patchy dune scrub and grassland vegetation including gaps, sparse

vegetation patches, erosional areas, and extensive low herbaceous vegetation in

varying successional stages indicate favorable conditions for colonization or

persistence by San Francisco lessingia. Population sizes discussed here refer to the

number (census or estimated population size) of mature, spontaneously established

seed-producing individuals of San Francisco lessingia, but not directly sown or

transplanted individuals, seedlings, or pre-reproductive plants.

Interim population criteria for this annual species are not based on management for or

maintenance of a fixed population size over time. The population criteria anticipate

large natural fluctuations over precipitation cycles. Precipitation cycles vary in length,

but are expected to be 5 to 10 years in duration. Prescribed population sizes are

expressed as minimum lower thresholds and expected upper targets. They do not

represent averages over multiple years. A general interim recovery criterion for all

sites is that each must be secured under long-term protection and management

favoring persistence (as described in the narrative outline). These recovery criteria

primarily address listing factors 1 (present or threatened destruction, modification, or

curtailment of its habitat or range) and 4 (the inadequacy of existing regulatory

mechanisms), and secondarily address listing factor 5 (other natural or manmade

factors affecting its continued existence).
128

i. Presidio Recovery Unit (Figures 10 and 11)

(1) Lobos Dunes Reserve. The restored Lobos Dunes site is now effectively

the core population for the species (Figure 11). Acceptable population size for

San Francisco lessingia here may fluctuate between 50,000 and more than

500,000 plants within a precipitation cycle. It would not be acceptable to

stabilize the population at or near 50,000 in a small portion of the site: it is

essential that lessingia numbers fluctuate, with occasional years of high

numbers, and that the species be distributed over most of the site. At this core

population, no significant net long-term decline in population size (e.g.,

decline between successive precipitation cycles) should occur relative to target

numbers. “Fluctuation” does not include changes in population size caused by

progressive changes in vegetation composition and structure that would occur

over decadal time scales (e.g., shifts in dominance to closed-canopy shrub

vegetation). 

In addition, interim recovery criteria for Lobos Dunes require that nonnative

vegetation (primarily annual grasses, but also iceplants [Carpobrotus sp.],

horseweed [Conyza sp.], Bermuda-sorrel, Cape ivy [Delairia odorata], radish

[Raphanus sativus] and sheep sorrel [Rumex acetosella]) be reduced to less

than 5 percent maximum cumulative annual cover (the peak cover percentage

of all invasive species, with different seasons for peak abundance of various

species), with progressive incremental decreases annually. Experience with

new methods may prove that somewhat higher or lower levels of invasive

species as proportions of total cover and plant cover may be the most efficient

means to maintain dune scrub habitat, in which case the new information

would provide the basis for future management. While it is a goal to locally

eradicate these invasive nonnative plants, it is a necessity to at least reduce

them to minimal levels. 

The management goal for recovery in the Presidio Recovery Unit is to achieve

a self-sustaining San Francisco lessingia population, continuous across the

recovery unit, by allowing the current populations to spread and join together. 

It is to be self-sustaining without intensive management, particularly a

dependence on large amounts of hand weeding. By establishing a larger area

of continuous dune scrub vegetation to reduce edge effects, together with 

Presidio

Golf Course

Site

Rob Hill

Site

Wherry

Dunes

Site

Lobos

Dunes Site

Public

Health

Services

Hospital

Site

Battery Caulfield

Road Site

RCD

RBL

P/E P/E

Baker

Beach

RCD

P/E

Rob Hill

P/E*

P/E

RCD

Wherry Housing RCDSL

P/E

RCD

P/E*

P/E

RCD

Mountain

Lake

Public Health

Services Hospital

RCD

Hospital Buffer

Rob Hill

Buffer

West

Washington

Housing

Building 1750

and Bowley Street

Interim Management Area

Special

Management

Zone

Special

Management

Zone

Lobos Creek

Upper Baker

Beach Dunes

Recovery Plan for

Coastal Plants of the

San Francisco Peninsula

Presidio Recovery Unit

Figure 11. Area Detail: Proposed reintroduction,

restoration, and protection areas within the

Presidio (Presidio Trust and Golden Gate National

Recreation Area) for recovery of San Francisco

Lessingia (Lessingia germanorum)

Lessingia germanorum

*

BL

CD

CDSL

P/E - Protect and enhance existing coastal

dune habitat

P/E - Restoration previously initiated (before

1993 photo); protect and ehance

restored habitat

R - Restore degraded coastal bluff

vegetation

R - Restore coastal dune grassland and

scrub vegetation suitable for reestablishment or expansion of

populations

R - Restore coastal dune slack vegetation

SMZ - Special Management Zone

Area requiring site specific planning

MAP PROJECTION: Transverse Mercator, UTM

Coordinate System, Zone 10, Datum NAD83

PHOTO SOURCE: Digital Orthophoto Quarter

Quads (DOQQ), USGS Images, June and July 1993,

One Meter Resolution.

1:2,000

GIS DATA: Digitized On-Screen

with DOQQs as backdrops by

USFWS, Sacramento Ecological

Services Office, GIS Branch

The Presidio Recovery Unit: a detailed look at the conceptual

design of a mosaic of restored and protected habitats, fitted to

the actual and potential habitat remnants of the modern

landscape. The configuration of the reserve design is based on

the distribution of surviving and rehabilitated populations of the

endangered San Francisco lessingia, and the topographic and

ecological relationships needed to link them and sustain them.

Ecological factors such as wind-dispersal of seed, wind-stress

effects on vegetation structure, slope, elevation, and aspect

(compass direction, in relation to exposure to sun and wind) are

important considerations for the design of the reserve.

0 0.04 0.08 0.16 0.24 0.32

Kilometers

Special Management Zone

Building 1750 and Bowley Street

Interim Management Area

West Washington Housing

to be evaluated

Boundary of Designated Area

Boundary of Buffer Areas

adjacent to Recovery Unit
131

continued work to develop improved management methods, we seek to

maintain a level of less than 5 percent weed cover of total cover. Current

methods, using intensive hand weeding, with current size of dune scrub habitat,

have not achieved weed cover to less than 15 percent of total cover.

To achieve recovery and enable delisting, we want to provide a satisfactory

level of assurance that ordinary, non-intensive management can be expected to

maintain site conditions. If we establish a non-intensive management regime,

and our monitoring data shows a thriving population of San Francisco lessingia

over the full range of weather conditions from wettest to driest precipitation

year types, with a controlled weed presence, we may be able to change the

management target to reflect the observed weed cover level. Until such time as

data shows thriving San Francisco lessingia over the entire range of annual

precipitation amounts in the presence of greater, but controlled, levels of

weeds, we will seek to find ways to meet the 5 percent target through adaptive

management.

The percentage of sand surface with bare/sparse dune surface (bare to sparsely

vegetated or dominated only by native annuals) must be at least 20 percent

cover at relatively large spatial scales (patch sizes greater than 3 meters [10

feet]) over the entire Lobos Dune site. Substrate conditions within 30

centimeters (12 inches) surface depth must match old dune sand (closely

similar texture, nutrient, and moisture holding capacity) and be conducive to

ecological regeneration of target dune vegetation.

Reference conditions for substrate conditions may be based on relict

population sites (e.g., Presidio Golf Course roadside, Rob Hill, behind Public

Health Services Hospital) where dune soils are not amended with imported fill

or augmented organic matter from nonnative vegetation. Non-dune sand

substrate that facilitates weed invasion or accelerated succession to mature

coastal scrub must be minimized to less than 5 percent of the managed dune

surface area of the Lobos dunes site.

(2) Battery Caulfield Roadside Reserve. This population (Figure 11) should

not decline below 1,000 plants in any three consecutive years. Population size

should intermittently reach or exceed 5,000. At this smaller, sheltered, partly
132

shaded site, nonnative vegetation (primarily annual grasses, but also iceplants)

must be reduced to less than 20 percent peak annual cover, with progressive

incremental decreases annually. Local extirpation of these invasive nonnative

plants is a goal, but reduction to minimal levels is necessary. The percentage

of sand surface with bare/sparse dune surface (bare to sparsely vegetated or

dominated only by native annuals) must be at least 10 percent cover.

(3) Wherry (Feral) Dunes Reserve. The current size of this site is

approximately 1 hectare (2.5 acres); it is expected to be enlarged (Figure 11). 

The population should not decline below 5,000 plants in any 3 consecutive

years. General vegetation criteria for Lobos dunes apply here.

(4) Rob Hill Reserve. The population should not decline below 5,000 in any 3

consecutive years, with intermittent years reaching or exceeding 10,000 over a

precipitation cycle (Figure 11). Vegetation criteria for the Battery Caulfield

site apply here.

(5) Public Health Services Hospital Reserve. This reserve includes the

Presidio Golf Course roadside site (Figure 11). The combined population size

of the Public Health Services Hospital site and Golf Course site should not

decline below 5,000 plants in any 3 consecutive years. Neither population

alone may decline below 1,000 plants in any year. Vegetation criteria for the

Battery Caulfield site apply here.

ii. Daly City Reserve. The population here should exhibit no net long-term

(progressive trend exceeding 3 years) decreases in spatial distribution or the

order of magnitude of population size within this site (Figure 6). The

population should not decline below approximately 50,000 plants over any 3

consecutive years. Intermittent peak year population sizes should exceed

approximately 200,000. The abundance (density and cover) and distribution of

nonnative vegetation (particularly iceplant and ripgut brome [Bromus

diandrus]) should exhibit no progressive (more than 2 consecutive years)

increases within any portion of the site that is persistently or intermittently

occupied by San Francisco lessingia. 
133

iii. Fort Funston Reserve. Establish recreation management needs for

maintenance of a dune scrub community in the Fort Funston Reserve through

completion of the site-specific studies identified in the Stepdown Narrative and

Implementation Schedule. A reintroduced population should be established at

Fort Funston and should persist over a full precipitation cycle. Precipitation in

California usually varies through most of the historical range from driest to

wettest during most 10-year periods. The reintroduced Fort Funston

population, derived from founders obtained from the Daly City population,

must achieve a minimum, self-sustaining size of approximately 500,000 plants

within 10 years after founders are transplanted.

iv. Offsite. Long-term seed storage will ensure that founder populations from

each site could be reestablished in the event of local extinction. Seed samples

totaling at least 500 seed from at least 25 separate parents (not clusters of

siblings in seedheads) sampled randomly throughout each site in its entirety

should be collected annually. Each annual collection should be placed in dry,

above-freezing refrigerated conditions at either the Golden Gate National

Recreation Area native plant nursery or local botanical gardens equipped for

long-term seed storage. Seed should also be deposited at a botanical garden

approved by the Center for Plant Conservation.

b. Long-term Recovery Criteria

All long term recovery criteria presume that the interim recovery criteria are

completely met. Long term criteria apply to the three recovery units: The Presidio

Recovery Unit should consist of dune complexes around nuclei of preserved sites,

many of which should eventually be connected and integrated in larger reserves. The

Southern Recovery Unit should consist of Daly City and Fort Funston reserves. The

Fort Funston Reserve should consist of restored dunes with reintroduced populations

of San Francisco lessingia from Daly City. The Fort Funston Reserve will include

restoration with native dune scrub species, and will require site-specific studies of

recreational use management consistent with maintaining restored dune scrub habitat

and recovery of San Francisco lessingia. The Daly City Reserve should consist of a

small area of managed vegetation including the preserved population there,

surrounded by native buffer vegetation zones in the same parkside and residential

setting. The Satellite Recovery Unit should consist of smaller reserves with San
134

Francisco lessingia populations relatively isolated from the main reserves in the

Presidio, Fort Funston, and Daly City. These reserves should be located on small open

space areas in San Francisco containing restorable dune remnants at Sutro Heights and

Sunset Heights. These recovery criteria primarily address listing criterion #1 (present

or threatened destruction, modification, or curtailment of its habitat or range), and

secondarily address listing criterion #5 (other natural or manmade factors affecting its

continued existence). For all reserves, buffers of landscape vegetation intended to

reduce conflicts, such as between building maintenance and management of San

Francisco lessingia, will exist around all historic structures. Vegetation management

for these buffers will include weed suppression or eradication.

The proposed recovery units must be permanently protected as natural area reserves

with vegetation management priorities and objectives dedicated to the persistence of

the San Francisco lessingia populations within the restored native dune vegetation and

dynamics that sustain them. The dedicated areas within lands of Golden Gate National

Recreation Area, Presidio Trust, and the City of San Francisco must be permanently

established, unaffected by potential land ownership transfers or proposed changes in

land use. Other park uses that are potentially compatible with the persistence of San

Francisco lessingia populations, such as well-regulated public access, may be retained

or modified to ensure compatibility. Incompatible park uses (planted trees or dense

shrub cover, road construction, irrigation, fertilizer application, construction of

buildings, etc.) must be excluded from the recovery units in the dune scrub habitat

following restoration, in perpetuity.

i. Presidio Recovery Unit 

Long-term recovery criteria apply principally to future populations of San

Francisco lessingia that have expanded into integrated, consolidated reserves

of restored dune areas around core remnant populations, or around large

founder populations derived from them. The long-term criteria, like the

interim criteria, are focused on specific geographic areas and their habitat

qualities. For long-term recovery, vegetation dynamics involving partial dune

instability (local blowouts with cycles of erosion, deposition, and stabilization)

are an essential qualitative criterion for the Lobos-Wherry-Baker and Fort

Funston reserves. The sizes and local distributions of San Francisco lessingia

populations in these reserves are expected to fluctuate substantially between
135

years. The criteria do not allow for artificially maintained large populations, as

could be achieved in the short term by intensive habitat manipulation (e.g.,

intensive manual weeding, re-seeding, artificial maintenance of early

successional stages of dune scrub). Instead, the criteria follow an ecosystem

management approach: they are based on natural fluctuations of population

size over large, heterogeneous areas of mature, high quality habitat. Numeric

population criteria are therefore only approximate, and magnitude of

population sizes relative to annual fluctuations is more pertinent than shortterm census data. 

The aim of long-term criteria is to ensure that resilient, dynamic landscape

units continue to support mobile populations of San Francisco lessingia that are

well-distributed in reserves that faithfully represent the native ecosystem of the

species. Failure of long-term recovery criteria would be indicated by

progressive declines of San Francisco lessingia populations, by progressive

deterioration of vegetation (excessive total vegetative cover and dune stability,

progressive increases in cover of invasive nonnative vegetation),

impoverishment of native species diversity within the reserves, or

establishment of fragmented or undersized reserves. Excessive or insufficient

vegetation management, such that natural vegetation and population processes

are overridden, would also indicate failure of recovery criteria. Overmanagement would be indicated by need for periodic “restocking”

(augmentation) of San Francisco lessingia populations by supplemental

seeding, or periodic replanting of native vegetation in previously restored dune

areas.

(1) Lobos-Wherry-Baker Dune Reserve. This section of the Presidio Recovery

Unit would incorporate and coalesce areas at Baker Beach dunes, Lobos

Dunes, Wherry Dunes and Housing sites, conifer groves around Lobos Dunes,

and the Battery Caulfield Road site (Figure 11). Criteria for this complex are

designed to ensure unobstructed wind fetch (from the Golden Gate, through

Baker Beach and its restored climbing dunes) and solar exposure. The

combination of large open fetch, high and steep slopes and extensive dune area

(approximately 44 hectares [110 acres]) here must sustain dune dynamics

favorable for patchy, variable dune vegetation and persistence of San Francisco

lessingia, minimizing the need for intervention to maintain specific phases of
136

dune succession. This complex must be large enough to enable turnover

(dynamic local extinction and colonization) of San Francisco lessingia patches,

and minimize the likelihood of population extinction. The reserve design need

not preclude other park uses and management of the reserve that are

compatible with these criteria.

(a) Structural habitat criteria. This complex must be restored to be a

continuous dune field. It must consist of a matrix of coastal dune scrub

and grassland, with a mosaic of active and stabilizing dune blowouts at

varying stages of local succession (erosion, deposition, and

stabilization phases). The dune complex must have: (i) effectively

unobstructed wind fetch to the Golden Gate, (ii) locally steep dune

slopes to ensure minimum persistent small-scale erosion and sparse

vegetation peripheral to blowouts and within stabilizing blowouts, and

(iii) full solar exposure. Progressive succession to stable, closed dune

scrub or grassland vegetation would conflict with this criterion. Largescale, intensive dune mobility (accretion/erosion greater than 25

centimeters [10 inches] per year in areas exceeding 0.2 hectare [0.5

acre]) may not occur over more than 15 percent of the complex in any

year. The portion of the reserve area with cover of either bare sand

(exclusive of primary pedestrian trails) or sparsely-vegetated sand

dominated by annuals and short-lived perennials should be a minimum

of 5 percent. These criteria should be achieved incrementally in phases

(involving demolition of buildings and removal of nonnative tree

plantings), at or near completion within 30 years.

(b) Population and vegetation criteria. As the size of the restored area

grows and improved weed control methods keep weed cover at minimal

levels inside the margins of the dune scrub habitat, our criteria for

measuring success are for nonnative vegetative cover to be maintained

below 5 percent (annual peak cover), with a declining trend at

locations in the restored area as further restoration pushes the margin

farther away. Experience with new methods may show that somewhat

higher or lower levels of invasive species as proportions of total cover

and plant cover may be the most efficient means to maintain dune scrub

habitat, in which case the new information would provide the basis for
137

future management. Population size of San Francisco lessingia within

this complex may exhibit extreme fluctuations locally (including local

extinction of colonies), but population size should not fall below

approximately 500,000 in any 3 consecutive years, and should be

distributed patchily (not continuously) over most or all of the complex. 

No maximum population size is set as a criterion, but peak population

size should intermittently exceed one to several million plants. Large

population size, per se, is not a preeminent criterion for long-term

recovery. Extremely high population sizes may be expected in early

successional phases of dune restoration sites, but are not likely to be

maintained when woody species achieve stable dominance patterns. 

Multiple new colonies (variable size patches of relatively high San

Francisco lessingia density) must become established spontaneously

(without artificial dispersal or supplemental seeding) in the complex

within 10 years. Population peaks of several million plants should

occur within 10 years. Criteria should be met in this reserve between

10 and 20 years (and no later than 30 years) after basic restoration work

(tree removal, structural demolition, regrading) has been completed in

the reserve.

Success at maintaining low levels of weeds and a self-sustaining San

Francisco lessingia population will: (1) keep weed seedbanks (amount

of seed in the top layer of soil) from irrupting and rising to levels where

they would suppress lessingia reproduction; and (2) keep lessingia

seedbanks from declining.

(2) Rob Hill Reserve (expanded; Figure 11) 

(a) Structural habitat criteria. The area supporting native dune

vegetation compatible with colonization and ecological regeneration of

San Francisco lessingia must be increased to approximately 2 hectares

(5 acres) by removing peripheral nonnative trees and groundcover

vegetation (blue gum [Eucalyptus globulus], iceplant, ivy, acacia

[Acacia sp.]). In addition, the southwest slope of Rob Hill facing the

expanded Wherry Dune Reserve must be restored to relatively sparse,

low dune scrub vegetation to increase connectivity between these
138

populations (low but biologically significant immigration of San

Francisco lessingia from the Lobos-Wherry-Baker Reserve) (Figure

11). Any expansion of this population will need to be coordinated with

trail and campground use. As noted above, buffers of landscape

vegetation of 15 to 30 meters (50 to 100 feet) around structures in the

Presidio would minimize the likelihood of management conflicts due

to colonization by San Francisco lessingia. However, historic

earthworks associated with batteries at this site may be compatible with

maintaining San Francisco lessingia populations. Site-specific planning

should consider reductions of the proposed buffer for this reserve. 

(b) Population and vegetation criteria. Population size should not fall

below approximately 100,000 (following expansion of habitat) at this

site for any 3 consecutive years, and San Francisco lessingia should be

distributed over most of the site. Population size should significantly

exceed this threshold in intermittent years, but no numeric peak targets

are specified. Peak population size would be expected to exceed one

million plants, particularly early in succession following dune

restoration. Nonnative vegetation (primarily annual grasses at this

location) must be reduced to less than 5 percent peak seasonal cover,

with progressive incremental decreases annually, aimed at local

extirpation. Experience with new methods may prove that somewhat

higher or lower levels of invasive species as proportions of total cover

and plant cover may be the most efficient means to maintain dune scrub

habitat, in which case the new information would provide the basis for

future management. Iceplant recolonization must be prevented by

recurrent removal of seedlings. If population size criteria are not met,

management must include periodic localized mechanical disturbance of

vegetative cover to expose patches of bare or sparsely vegetated dune

sand on approximately 2 to 5 percent, and up to 15 percent, of the

restored dune area at least once every 7 years, or more often. 

Vegetation trends over a period of 20 years must exhibit long-term

persistence of large areas dominated by annual native vegetation or

sparse cover of mixed native annuals, perennials, and shrubs, over at

least 25 percent of the site.
139

(3) Public Health Services Hospital Site (expanded; Figure 11)

(a) Structural habitat criteria. The vegetation on the slope

behind (north of) the Public Health Services Hospital should be

modified to promote the necessary southwest wind influence,

promoting viable San Francisco lessingia habitat on the upper

plateau of that slope (Figure 11). A vegetated buffer of low

statured (less than 6 meters in height), non-invasive native

plantings may be established to prevent any potential conflict

with building operations that could result in any inadvertent

degradation of San Francisco lessingia habitat on the upper

plateau of that slope.

The restored dune habitat occupied by patches of San Francisco

lessingia must be at least 3 hectares (7 acres). Nonnative trees

and shrubs around the local oak woodland dune remnant must

be removed from the areas to be restored to dune scrub habitat,

but the remnant native dune vegetation should not be subjected

to any earthwork. Topographic relief of regraded slopes must

be irregular, including steep local slopes with variable aspects. 

Management must include periodic mechanical disturbance of

restored (not remnant woody) vegetative cover to expose

patches of bare or sparsely vegetated dune sand on

approximately 2 to 5 percent (but up to 15 percent) of the

restored dune area at least every 7 years, or more often if

population criteria are not met.

(b) Population and vegetation criteria. The local San Francisco

lessingia subpopulation must colonize the cleared plateau at the top of

the slope behind the Public Health Services Hospital (former Marine

Hospital site). Population size in the remnant and restored areas should

intermittently reach or exceed approximately 300,000 within 10 years,

and should not fall below approximately 50,000 in any 3 consecutive

years. In addition, nonnative herbaceous vegetation (primarily annual

grasses) should be reduced to less than 5 percent peak seasonal cover,

with progressive incremental decreases annually, aimed at local
140

extirpation. Experience with new methods may prove that somewhat

higher or lower levels of invasive species as proportions of total cover

and plant cover may be the most efficient means to maintain dune scrub

habitat, in which case the new information would provide the basis for

future management. Woody nonnative vegetation must be prevented

from vegetationally regenerating after removal. Vegetation trends over

a period of 20 years must exhibit long-term persistence of large areas

dominated by annual native vegetation or sparse cover of mixed native

annuals, perennials, and shrubs, over at least 25 percent of the site.

ii. Southern Recovery Unit

(1) Daly City Reserve (expanded)

(a) Structural habitat criteria. Because this site is not part of a dune system,

and was artificially created by grading activities, structural criteria do not

apply. Vegetation management by humans, rather than reestablishment of

natural dune ecosystem processes, will be applied in this reserve.

(b) Population and vegetation criteria. At least 1.2 hectares (three acres) of

this site must be (gradually) cleared of nonnative vegetation, to establish a

minimum core population of at least 50,000 plants under actively managed

conditions (Figure 6). The overall population should exhibit no net long-term

(progressive trend exceeding 3 years) decreases in spatial distribution or order

of magnitude of population size within this site. Minimum population size

(combined for the intensively managed core area and areas beyond it) for any 3

consecutive years is approximately 50,000 plants, with intermittent peak years

exceeding approximately 200,000 (the same as for the interim criteria). If

population size criteria are not met, management should include periodic

mechanical disturbance of restored (not remnant woody) vegetative cover to

expose patches of bare or sparsely vegetated sand on approximately 2 to 5

percent, and up to 15 percent, of the managed sandy coastal scrub area at least

every 7 years. The abundance (density and cover) and distribution of

nonnative vegetation (particularly iceplant and ripgut bromegrass) should

exhibit no significant increases within any portion of the site that is persistently

or intermittently occupied by San Francisco lessingia. 
141

(2) Fort Funston Reserve (reintroduction) (Figure 12):

(a) Structural habitat criteria. A total of approximately 30 hectares (75 

acres) of the reserve, including at least 26 hectares (65 acres) of 

continuous restored dune area at the north end of Fort Funston (north end

reserve), should be dedicated to dune restoration and vegetation

management favorable to San Francisco lessingia sufficient to maintain

ecosystem functions and support a self-sustaining population of San Francisco

lessingia (Figure 12). This reserve will also include management buffers

around all historic structures. Plantings would be native dune scrub species

and vegetation management would include weed suppression. Recreational

use management of restored dune scrub on this reserve will be determined

through site-specific studies. An action for this is included in the Stepdown

Narrative.

In addition, a 4-hectare (10-acre) area of remnant dunes near the southern end

of the Fort Funston Reserve, potentially subject to mixed use, should be

managed as a subsidiary area for a reintroduced San Francisco lessingia

population. 

The desired recovery objective for the 26-hectare (65-acre) north end reserve is

to restore habitat composed of a matrix of low-growing coastal dune scrub and

grassland, with a mosaic of active and stabilizing dune blowouts at varying

stages of local succession (erosion, deposition, and stabilization phases). The

north end reserve should have unobstructed wind fetch to the sparsely

vegetated marine bluff, with no conifer, blue gum, or nonnative shrub cover

upwind or peripheral to the reserve. Stabilizing iceplant mats must be removed

from the reserve, including debris. The dune complex should have locally

steep dune slopes to ensure minimum persistent small-scale erosion and sparse

vegetation peripheral to blowouts, and within stabilizing blowouts. 

Progressive succession to stable, closed dune scrub or grassland vegetation

would conflict with this criterion. Large-scale, intensive dune mobility

(accretion/erosion greater than 25 centimeters [10 inches] per year in areas

exceeding 0.2 hectare [0.5 acre]) must not occur over more than 15 percent of

the north end reserve. Percentage of the reserve area with cover of either bare

sand (exclusive of primary pedestrian trails) or sparsely-vegetated sand 
P/E

Lake Merced

Lake Merced

M LG

R

CD

R CD

R

REC

Golden Gate

National Recreation

Area Boundary

Fort

Funston

Recovery Plan for

Coastal Plants of the

San Francisco Peninsula

Fort Funston

 and Vicinity

Figure 12: Southern Recovery Unit for

San Francisco Lessingia

Fort Funston proposed reintroduction,

restoration, and protected areas within

Fort Funston (Golden Gate National

Recreation Area) dunes, near historic

Lake Merced Collection Locality.

(Lessingia germanorum)

MAP PROJECTION: Transverse Mercator, UTM

Coordinate System, Zone 10, Datum NAD83

PHOTO SOURCE: Digital Orthophoto Quarter

Quads (DOQQ), USGS Images, June and July 1993,

One Meter Resolution.

0 0.09 0.18 Kilometers

1:3,132

M - Manage existing habitats compatible

with sustainable reestablished

populations of

P/E - Protect and enhance existing habitat

R - Restore coastal dune habitat

suitable for

restablish populations

R - Rehabilitate degraded coastal bluff

landslides for recreation-priority use

to relieve intensive recreational

pressures on Fort Funston dune

habitats

Lessingia germanorum

CD

REC

GIS DATA: Digitized On-Screen

with DOQQs as backdrops by

USFWS, Sacramento Ecological

Services Office, GIS Branch

LG

Lessingia germanorum

Boundary of Designated Area

Fort Funston is the main component of the

Southern Recovery Unit for San Francisco

Lessingia. It is the southwestern tip of the

San Francisco dune system, and overrides

raised ancient coastal sand deposits, the

Merced bluffs. This is a multiple use park,

with heavy recreational use, especially near

the trails close to the parking areas. San

Francisco lessingia may be quite compatible

with appropriate levels of disturbances and

vegetation gaps that trampling pressures

provide. To help distribute recreational uses

with conservation of native habitat, the recovery

plan proposes to rehabilitate an extensive area

south of the National Park boundary, and

dedicate it to recreation-priority uses. It also

proposes a gradient of management within the

National Park, ranging from no endangered

species management, mixed recreation and

experimental endangered plant habitat, and

restored coastal dune habitat with priority for

conservation of San Francisco lessingia and

associated native species.
143

dominated by annuals and short-lived perennials should be a minimum of 15

percent. Trampling impacts that cause partial devegetation of dunes, if rotated

among areas to alternate between years of disturbance and years of protected

revegetation, may be compatible with this criterion at this locality. This

criterion should be achieved incrementally in phases within 10 years.

The approximately 4-hectare (10-acre) south end reserve would be less

intensively managed or restored for San Francisco lessingia, and criteria would

be accordingly more flexible. The 4-hectare (10-acre) reserve will be located

within the southern portion of the Fort Funston Reserve. Vegetation

dominated by iceplant, Monterey cypress (Cupressus macrocarpa), or other

nonnative vegetation must not exceed 40 percent of the area. Cover of sparse,

low, native dune vegetation structurally suitable for San Francisco lessingia

must be at least 5 percent. Intensively trampled areas that effectively preclude

establishment of San Francisco lessingia seedlings may not exceed 60 percent

of the area.

(b) Population and vegetation criteria. The reintroduced Fort Funston

population, derived from founders obtained from the Daly City population,

must achieve a minimum size of approximately 500,000 plants within 10 years

after founders are transplanted. The use of the Daly City seed source is based

on genetic testing showing that population is genetically distinct from that of

the Presidio. Population size may be expected to reach millions of plants

temporarily in early phases of dune succession after restoration, but extremely

large population size is not prescribed as a recovery criterion. The population

must exhibit no progressive long-term (more than 3 consecutive years)

declines, and a minimum annual population size of 10,000 plants the fifth year

after reintroduction. Multiple new colonies must spontaneously establish

outside sites of introduction (with no additional artificial seeding) within the

complex within 10 years after implementation of dune restoration work. The

population should at least initially be distributed in discrete or coalescing

multiple colonies. Cover by nonnative woody vegetation within the reserve

must be below 1 percent at any time. Cover of iceplant and European

beachgrass by year 10 must be below 10 percent of the reserve area, and must

decline progressively. Iceplant must be effectively eradicated from the reserve

by year 20, and its density in adjacent buffer areas must be reduced to less than
144

10 percent. Native dune scrub and grassland vegetation with ample open,

partially and moderately disturbed patches should dominate this reserve area. 

iii. Satellite Recovery Unit

The Satellite Recovery Unit consists of reserves at Sunset Heights and the

Sutro Heights/Cliff House/Lands End area dune remnants. The specific size,

number, and configuration of reserves needs to be determined on the basis of

more detailed site-specific information than is currently available. 

The exact location of restorable dune habitat areas, and their relationship to

particular parcels and current ownership was not available during the

development of the recovery plan. We will work to find opportunities where

there is suitable habitat and willingness of owners to participate or willing

sellers and willing buyers who are interested in restoring dune scrub habitat.

Preliminary criteria for this unit are as follows: cumulative population size

among all of the remnant dune sites must reach 100,000 plants within 10 years

after founders are introduced, with a minimum cumulative annual population

size of 5,000 plants for at least 3 consecutive years. Population size criteria

should be revised for the Satellite Recovery Unit when the carrying capacity

and management constraints of these small sites are better understood through

experience and adaptive management. Criteria for vegetation management at

the southern reserve at Fort Funston apply to these sites, but management goals

should strive to exceed criteria.

iv. Areas adjacent to the Presidio Recovery Unit to be evaluated. The Presidio

Trust has proposed that restoration of dune scrub habitat to enlarge the Presidio

Recovery Unit following the removal of West Washington Housing over the

next 30 years may increase the likelihood of recovery for San Francisco

lessingia. The West Washington Housing area is located adjacent to both the

Wherry Housing and Rob Hill Reserves. The suitability of the site for addition

to the Presidio Recovery Unit and potential to expand the currently proposed

reserves should be evaluated and incorporated into the recovery plan, if

appropriate. We will evaluate the addition of the West Washington Housing

area during implementation of the recovery plan to determine whether it will
145

contribute to the continuous functional habitat necessary to reduce edge effects

on the San Francisco lessingia population and help achieve recovery.

v. Buffer areas adjacent to the Presidio Recovery Unit. Two buffer areas

adjacent to the Presidio Recovery Unit and to populations of San Francisco

lessingia are not suitable for restoration to dune scrub habitat but their

management can contribute to the long-term viability of San Francisco

lessingia populations by acting as buffers to reduce the threat of invasive

weeds. These two areas are currently proposed by the Presidio Vegetation

Management Plan to retain the current vegetation character, but to be managed

to reduce populations of invasive weeds. These areas include the Rob Hill

Buffer and the Public Health Services Hospital Buffer. 

The Rob Hill Buffer is located adjacent to the northeast edge of the Presidio

Recovery Unit (see Figure 11). At the northeast edge of the recovery unit, on

Rob Hill, is an area to the east of the coastal dunes. On the back slopes of this

area, there are areas of forest. In general, the buffer is to be managed

according to the Presidio Vegetation Management Plan. On the edge adjacent

to the dune scrub habitat, the trees will be managed to create a transition to the

edge that will not extend above the height of the topography of the dune scrub. 

The area is also to receive special attention for weed control, the chief edge

effect to be buffered against. Horticultural management use of supplemental

irrigation and fertilization is to be done to avoid increasing the water and

nutrient supply in the dune scrub area.

Public Health Services Hospital Buffer Area is located adjacent to the

southeast edge of the Presidio Recovery Unit (see Figure 11). At the southeast

edge of the recovery unit, above the Public Health Services Hospital, is an area

where the plateau of relictual dunes slopes steeply down to the hospital and

some housing just to the northeast of the hospital. The area is also to receive

special attention for weed control, the chief edge effect to be buffered against. 

Horticultural management use of supplemental irrigation and fertilization is to

be carried out in a manner to avoid increasing the water and nutrient supply in

the dune scrub area.
146

2. Recovery Criteria for Raven’s Manzanita

Recovery criteria for Raven’s manzanita apply to the single remaining wild plant and

the existing transplanted clones in the Presidio (interim criteria), to future expanded

populations in restored habitats within recovery units, and to conditions of the plant

communities in which they occur. Interim criteria are aimed at ensuring that the

species as a whole does not become extinct, and that the one remnant wild clone does

not die prematurely from threats. The eventual natural death of the wild clone is not

expected in the foreseeable future. Interim criteria are also aimed at ensuring that

cultivated populations of Raven’s manzanita are maintained, and that essential

information regarding its reproductive biology and potential for reintroduction is

investigated. Long-term recovery criteria for Raven’s manzanita are aimed at

establishment of multiple new sexually reproducing populations in local serpentine

vegetation reserves that approximately reassemble the local serpentine flora associates

of Raven’s manzanita at interior San Francisco localities. Long-term recovery also

aims to ensure survival of the remnant clone by increasing its distribution within

unoccupied suitable serpentine habitat in the Presidio. These criteria are also aimed at

ensuring that populations are not artificially perpetuated solely by vegetative

propagation of one clone. If possible, the species should genetically regenerate

appropriate variability in ecological, morphological, and genetic characteristics,

similar to that of other narrowly endemic manzanitas. Such variability is presumed to

be important for continued adaptation and evolution of Raven’s manzanita, as for most

sexual species (Stebbins 1950, Ghiselin 1988).

Raven’s manzanita may be considered for reclassification to threatened status when all

interim recovery criteria are fully achieved, and: (1) at least five spontaneously

reproducing variable populations are established in reserves on bedrock outcrops

outside the Presidio in San Francisco, at least three of which must be on serpentine

outcrops; (2) at least two sexually reproduced generations are established within the

Presidio; and (3) at all the sites, population size and individual clone size increase over

a period of 30 years. These criteria provide a trigger for a review of Raven's

manzanita's eligibility for downlisting from "endangered" to "threatened" status. 

They represent partial fulfillment of long-term recovery criteria, focusing on indicators

of sufficient progress towards the long-term recovery goals to justify reconsideration

of the level of protected status needed. Because recovery is constrained by the small

area of potential restorable habitat in San Francisco, reclassification to threatened
147

status is not foreseeable in less than 30 years, and may not be feasible. Furthermore,

recovery sufficient to warrant full delisting of Raven’s manzanita (removal of all legal

protection under the Endangered Species Act) is not projected within the foreseeable

future and may not be possible. This conclusion may be reconsidered in the remote

future if success of long-term recovery exceeds current expectations.

a. Interim Recovery Criteria

i. Habitat and Population Stabilization in the Presidio. The site of the

original remnant clone and all daughter clones established in the Presidio must

be dedicated to permanent habitat protection, maintained, and protected in

perpetuity (principally by removing nonnative vegetation). The original and

daughter clones must exhibit significant net growth (see Appendix IV) over a

10-year period, while the number of daughter clones must increase. These

recovery criteria address listing factor 1 (present or threatened destruction,

modification, or curtailment of its habitat or range) and listing factor 4 (the

inadequacy of existing regulatory mechanisms).

ii. Propagation of Seedling and Clonal Stock. Multiple nursery populations of

propagated Raven’s manzanita must be established within the Golden Gate

National Recreation Area and at two or more botanical gardens that are

committed to conservation of this species. Nursery populations must consist of

both clones and seedling-grown plants. Seedling-grown plants must be derived

at least from self-pollinated inbred lines (highest priority), but may include

separate experimental breeding lines composed of recurrent backcrosses of

selected Tamalpais manzanita (Arctostaphylos montana) individuals (and

possibly Franciscan or other manzanita taxa) on Raven’s manzanita if

production of inbred lines is not feasible, and if the strategy is recommended

by a scientific review panel of manzanita experts, plant conservation

geneticists, and others (see Recovery Strategy). The panel should develop a

genetic management plan in cooperation with us before any hybridization. 

Genetic management of the species should be subject to expert peer review. 

Artificially bred stock should be maintained in both permanent outdoor

collections for unrestricted growth (and future potential propagation stock),

and in container-grown collections available for outplanting at restoration sites. 

The total cultivated population size must be maintained at 50 or more daughter
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clones (of original Presidio plant) at all times, with a goal of 50 seedling plants

(preferably inbred, at least initially) that have at least two clonal replicates each

(total 200 plants). This recovery criterion primarily addresses listing factor 1

(present or threatened destruction, modification, or curtailment of its habitat or

range) and listing factor 5 (other natural or manmade factors affecting its

continued existence).

iii. Establishment of New Daughter Clones on Presidio Serpentine Bluff Sites. 

At least five additional colonies, each comprising at least five of the daughter

clones (with a goal of at least five inbred seedling-grown plants), must be

established on relatively stable, exposed serpentine outcrops within or above

the Presidio bluffs or in suitable inland outcrop areas, in areas where preexisting vegetation is sparse, particularly on steep slopes. New colonies must

exhibit net growth 5 years after transplanting with intensive maintenance, and

for an additional 5 years after cessation of intensive maintenance. This

recovery criterion primarily addresses listing factor 1 (present or threatened

destruction, modification, or curtailment of its habitat or range).

iv. Investigation of Taxonomic Relationships and Reproduction. Studies must

be conducted to clarify the taxonomic relationships between Raven’s

manzanita and Monterey County Arctostaphylos hookeri subspecies

hearstiorum and hookeri, Tamalpais manzanita (A. montana), bearberry

(A. uva-ursi), Franciscan manzanita (A. franciscana), and other relevant taxa. 

The breeding systems of these taxa, including comparisons of fruit set and seed

viability resulting from within-species crosses and self-pollination, should also

be studied. An especially high priority is to experimentally determine the level

of self-compatibility (level of viable seed production resulting from selfpollination) in the one remaining clone of Raven’s manzanita. Other topics to

investigate include variation in reciprocal hybrids among these species, pollen

viability of interspecific (between-species) hybrid plants, and chromosome

counts of hybrids. Experimental studies of natural ecological conditions

favoring seed production and seedling recruitment should be conducted. 

Fitness of inbred Raven’s manzanita plants (e.g., comparison of characteristics

related to growth rate, plant size, and reproductive traits) , and reference

populations of Tamalpais manzanita from serpentine sites (or other appropriate
149

reference populations of representative manzanitas) should be studied on

serpentine and nonserpentine substrates.

Results of these investigations will make it possible to evaluate the need for an

introgressive breeding program to restore sexual reproduction and adaptive

variability in the species. If introgressive breeding of Raven’s manzanita is

justified by scientific evaluation of its reproductive biology and taxonomy (see

(b) above), similar studies would probably be needed on backcross breeding

lines as well. The need for such research should be addressed in the genetic

management plan. This recovery criterion primarily addresses listing factor 5

(other natural or manmade factors affecting its continued existence). 

b. Long-term Recovery Criteria

The long-term recovery criteria below represent the maximum level of recovery that

we reasonably expect at present, given the irreversible urban constraints to habitat

restoration and population expansion. Fulfillment of these criteria is important as a

management goal to minimize the threats to the species as much as possible, but is not

considered sufficient to warrant delisting. Delisting may be reconsidered in the distant

future if success of long-term recovery significantly exceeds current expectations.

i. Reproduction and Growth in the Presidio Bluff Population. If feasible, at

least one generation of spontaneously established inbred (not experimental

hybrid backcross) seedlings of Raven’s manzanita must grow to reproductive

maturity in at least one colony out of five new Presidio bluff subpopulations

within 30 years after establishment. Over 50 percent of plants within all five

colonies must exhibit progressive and significant net growth (see Appendix IV)

over 20 years. This recovery criterion primarily addresses listing factor 1

(present or threatened destruction, modification, or curtailment of its habitat or

range) and secondarily addresses listing factor 5 (other natural or manmade

factors affecting its continued existence). 

ii. Establishment and Protection of New Interior Populations. At least five

mixed populations (Franciscan and Raven’s manzanita) consisting of original

clones and cloned seedlings (preferably inbred lines, if they are feasible and

found to be suitable for reintroduction to novel reintroduction sites) must be
150

established at separate interior San Francisco serpentine outcrop sites. Over 50

percent of founder plants at each new population must exhibit net growth in

size over a 10-year period. At least one generation of spontaneously recruited

seedlings of Raven’s manzanita must establish within 25 years in at least one

interior site. Significant recurrent production of viable seed must be in

evidence at all five sites. All reintroduction sites must be permanently

protected and monitored, and must be permanently maintained to prevent

reinvasion by competing nonnative vegetation, degeneration from recreational

misuse, or unforseen threats that require adaptive management. This recovery

criterion primarily addresses listing factor 1 (present or threatened destruction,

modification, or curtailment of its habitat or range) and listing factor 4 (the

inadequacy of existing regulatory mechanisms). 

iii. Permanent Reserve Cultivated Populations in Botanical Gardens. 

Horticultural propagation of Raven’s manzanita (also interim recovery criteria)

must be dedicated in perpetuity at no fewer than four botanical gardens in

California. Multiple independent garden collections in different California

coastal regions reduce the chance that region-wide catastrophic events (e.g.,

virulent new pathogens, extreme rainfall) could cause general loss from

cultivation. Propagation and cultivation of Raven’s manzanita for other

specific educational, scientific, or outreach efforts in support of recovery

actions recommended in this plan may be needed on a case-by-case basis for

recovery implementation, but such propagation and cultivation are not treated

as recovery criteria. This recovery criterion primarily addresses listing factor 1

(present or threatened destruction, modification, or curtailment of its habitat or

range).

B. Comprehensive Strategy of Recovery Actions

1. Recovery Strategy for San Francisco Lessingia

Recovery actions are discussed below in the context of independent geographic units. 

Recovery actions must be implemented sequentially as described below in order to

function properly. For example, reintroduction of endangered species before adequate

site preparation is completed (control of invasive nonnative plants, soil contouring,

establishment of wind fetch corridors, buffers) is likely to fail or require excessive
151

remedial intervention. Similarly, piecemeal restoration of large sites or patches within

larger sites is likely to result in excessive edge effects that diminish the habitat value

of restored areas, and impede seed dispersal linkages among sites. These recovery

actions address both interim and long-term recovery criteria. They are presented as

the conceptual basis for subsequent management and restoration plans. 

a. Presidio Recovery Unit

i. General Requirements of the Presidio Recovery Unit. The basic strategy for

the Presidio Recovery Unit of San Francisco lessingia is twofold: (1) maintain

and improve habitat quality of existing populations by suppressing or

eradicating competing nonnative vegetation; and (2) enlarge existing

populations and habitat by completing phased, large-scale dune scrub and

grassland restoration. Dune restoration here places emphasis on

reestablishment of natural vegetation patterns, topography, exposure, and

dynamic disturbance regimes that will maintain suitable microenvironments for

San Francisco lessingia and associated dune annuals. Restored vegetation may

include elements of native trees and shrubs such as live oak and arroyo willow,

at densities and distributions similar to natural dune scrub communities.

The Presidio populations of San Francisco lessingia are on Federal lands

managed by the Golden Gate National Recreation Area (National Park

Service), and owned by either the Golden Gate National Recreation Area

(Lobos Dunes restoration site, now the largest discrete population) or the

Presidio Trust (Rob Hill, Golf Course, and Public Health Services Hospital

sites, Wherry Dunes restoration site at the northwest end of Wherry Housing). 

The geographic scope of the Presidio Recovery Unit for San Francisco

lessingia is limited by the distribution of large blocks of dune sand substrate in

areas that are covered by scrub, nonnative trees, or nonhistoric buildings

previously programmed for demolition. These blocks of substrate occur

mainly in the southwest corner of the Presidio. Developed lands (e.g.,

permanent buildings, golf course, roads) constrain the potential configuration

and area of restored dune. The existing San Francisco lessingia sites are

separated from each other by nonnative trees (pine, cypress, blue gum groves)

and developed areas with roads and buildings. A nonnative tree grove between

Lincoln Boulevard and Baker Beach also creates a wind barrier that obstructs
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the fetch of strong winds from the Golden Gate and Baker Beach, and

adversely shelter the native dune vegetation. A final negative impact of

nonnative trees is that they add nutrients, moisture and shade to adjacent San

Francisco lessingia habitat, making it less suitable. These sites must be

reunited to a contiguous block of dune habitat capable of dynamic internal

redistribution of sand and seeds. Surface contours of tree-cleared dunes must

be modified to include steep, hummocky slopes prone to small-scale erosion

that will sustain local vegetation gaps for San Francisco lessingia to colonize. 

Recognizing that many of the non-native trees are part of the historic forest

that contribute to the National Historic Landmark status of the Presidio,

adaptive management strategies will be required to provide optimal wind fetch

and other ecosystem function while attempting to retain the character of the

historic forest. Several recovery actions describe processes to determine

appropriate management of historic forest within several portions of the

Presidio Recovery Unit.

The first requirement for recovery of San Francisco lessingia in the Presidio, to

control invasive nonnative vegetation, is most important in short-term and

near-term survival of the species. The second requirement, to have large

blocks of restored dunes with appropriate slopes, orientation, substrate,

mobility and wind-fetch, is essential to long-term recovery of the species. 

Dune restoration must occur at a spatial scale and setting that will allow for

blowout dynamics; otherwise, natural vegetation succession toward dense dune

scrub would over time reduce or eliminate open microhabitat for San Francisco

lessingia. Dune restoration will require extensive removal of nonnative blue

gum and conifer groves (e.g., Figure 11). The Lobos-Wherry-Baker lessingia

reserve will also require phased demolition of the Wherry Housing area, which

was programmed in the Presidio General Management Plan

Amendment/Environmental Impact Statement (National Park Service 1994)

and the Presidio Trust Management Plan (Presidio Trust 2002). Restoration of

a natural, dynamic regime of ecological disturbances and stress maintained by

interactions of wind, slope, and dune sand is essential to maintain conditions

that will support San Francisco lessingia and suppress invasive nonnative

species. Because the Presidio Recovery Unit overlaps sections of the historic

forest that contribute to the National Historic Landmark status of the Presidio,

major visual landscape changes such as tree removal in this highly visited part
153

of the park, adjacent to residential areas, will require substantial public

outreach and education for support. Many of the non-native trees that are of

concern in providing for the recovery of the San Francisco lessingia are part of

that forest. Adaptive management strategies will be developed to provide

optimal recovery conditions while attempting to retain the overall historic

character of the forest. In cooperation with the Presidio Trust and the National

Park Service, iterative strategies of thinning, limbing, increasing space

between trees, and even change out of species to lower growing but historically

consistent trees may be tested to increase ecosystem function, such as wind

fetch. Monitoring of the effectiveness of these strategies and revising them as

necessary to meet recovery goals will be essential to the recovery process.

While substantial progress at control of invasive nonnative plants has been

achieved at managed San Francisco lessingia sites of the Presidio, reinvasion

pressures remain strong, sustaining the need for intensive manual weeding

each year. Six years of monitoring data indicate that even with several

thousand hours of weeding done every year, the cover of nonnative plants is

greater than 12 percent for both the Lobos Dunes and Feral Dunes restoration

sites (Brian O’Neill, National Park Service, in litt. 2002). Currently, staff,

volunteer, and conservation corps labor through the Presidio Park Stewards

Program is the mainstay of weeding. Iceplants and annual grasses, especially

brome, have recently been principal weeds of the Presidio sites. Expansion of

restored and managed dune vegetation will probably exceed the capacity of

Presidio Park Stewards weeding at its foreseeable levels. Because native

grasses occupy a small proportion of these lessingia sites (mostly through

reintroduction in 1996), weed management should be expanded to include

experimental applications of appropriate grass-specific herbicides within and

around San Francisco lessingia sites (including adjacent source population

areas). The experimental program should continue over at least several years,

with the aim to exhaust local seed banks of nonnative invasive grasses. 

Manual weeding would be more effective after reduction of population density

and seed bank size of invasive annual grasses. Increased volunteer

participation should be encouraged, but we recommend that resources for

trained park vegetation management technicians be ensured so that weed

management can be performed at critical times of the growing season

(especially pre-flowering and during seed set) regardless of volunteer levels.

We also recommend that the measures needed to meet weed management
154

objectives, and the staffing to carry them out (whether by volunteers led by

staff, or by staff alone), be analyzed in a nonnative vegetation control plan for

all Presidio sites that support San Francisco lessingia and the adjacent buffer

areas.

ii. Site-specific Requirements of the Presidio Recovery Unit. The following

recovery requirements are presented as preliminary conceptual restoration and

reintroduction plans. Conceptual plans must be adapted to site-specific current

conditions. Site-specific plans must incorporate detailed information on soils,

topography, schedules, costs, engineering and construction methods and

requirements, etc. 

(1) Lobos Dunes Reserve. The Lobos Dunes restoration site must expand

further into what are currently degenerating groves of decadent Monterey

cypress trees extending to Battery Caulfield Road and Lincoln Boulevard

(Figure 11). Expansion of this core population area is the highest restoration

priority for the species, and should precede, or occur concurrently with, dune

restoration at any other lessingia reserves, as appropriate environmental review

is completed. The majority of this site is within the Special Management Zone

identified in the Presidio Vegetation Management Plan. Adaptive management

strategies will be developed to provide optimal recovery conditions while

attempting to retain the overall historic character of the forest where it overlaps

with the Presidio Recovery Unit. Tree removal must include removal of

woody debris, most stumps, and duff. Restoration actions should include at

least 2 years of substrate preparation and weed eradication. During this time,

revegetation with native species (including San Francisco lessingia) may be

appropriate. Substrate preparation for restoration at this location should

include measures to probe for significant seed banks of native species, and to

exhaust invasive nonnative weed seed banks. Surface recontouring following

seed bank probes must establish appropriate dune topography, including

oversteepened northwest-facing erosion-prone slopes. Exposed, bare dune

surfaces will need to be temporarily stabilized with inert materials (such as

desiccated, dead beachgrass) prior to revegetation. A buffer area of more

continuous stable dune scrub vegetation may be maintained along Battery

Caulfield Road, to limit potential sand movement conflicts with road

maintenance. Expansion of the Lobos Dunes restoration site will decrease

adverse edge effects (shading, fog drip, wind sheltering, leaf litter and leachate,
155

and weed reinvasion) on the existing Battery Caulfield lessingia site. 

Expansion of habitat will also reduce the risk that San Francisco lessingia

populations will significantly decline due to excessive vegetative cover and

over-stabilization of dunes.

(2) Wherry Dunes and Baker Beach. The Wherry Housing area above Baker

Beach is contiguous with the Lobos Dune site (Figure 11). It has the best

potential for restoration of dynamic dune scrub and blowout habitat on the

Presidio because of its elevation, aspect (facing northwest winds from the

Golden Gate, with potentially unobstructed fetch), steep slopes, and underlying

old dune sands. The 1-hectare (2.5-acre) Wherry (Feral) Dunes Restoration

site at the north end of Pershing Drive should be expanded both north (to

Washington Boulevard) and east (towards Lobos Dunes) in phases, eventually

coalescing with the Lobos Dunes restoration site. This expansion will require

phased demolition of buildings and removal of the nonnative pine and blue

gum stands around Pershing Drive and Washington Boulevard, so that old

dune scrub above Baker Beach and remnant dune scrub northeast of Wherry

Housing area (“Feral Scrub”) become incorporated in the complex. A portion

of this site is within the Special Management Zone identified in the Presidio

Vegetation Management Plan. Adaptive management strategies will be

developed to provide optimal recovery conditions while attempting to retain

the overall historic character of the forest where it overlaps with the Presidio

Recovery Unit.

The proposed Wherry-Lobos Dunes restoration complex would be adversely

affected by the strong wind-sheltering effects of the planted conifer and blue

gum groves between Lincoln Boulevard and Baker Beach unless these groves

are removed or modified. These trees obstruct the wind-fetch from the Golden

Gate (Figure 11). The wind-shadow of these tree groves would inhibit windshearing of prostrate vegetation and reduce the initiation and development of

small dune blowouts in the restored dune area. Wind-shear stress, drying and

post-blowout revegetation are the natural process most likely to maintain

suitable microhabitats for San Francisco lessingia. Nonnative trees also

directly displace suitable habitat for San Francisco lessingia and associated

species of concern in the dunes above Baker Beach, southwest of the “sand

ladder” (pedestrian walkway) on the steep dune slope at the northeast end of

Baker Beach. Most or all of these trees must be removed or forest
156

management practices modified to restore the wind fetch that is essential for

the long-term viability of the Wherry-Lobos Dunes. Interim forest

management practices (replacement with lower stature trees, lower tree

densities, higher tree canopy floor, etc.) described in the Presidio Vegetation

Management Plan may be implemented to improve ecosystem function,

including wind scouring, weed control, moisture balance, and San Francisco

lessingia recruitment. However, if aspects of ecosystem function are not

adequate to meet recovery objectives, additional measures (and potentially

complete removal of trees) to improve ecosystem function should be developed

in coordination with the Presidio Trust and Golden Gate National Recreation

Area. The trees should be removed or ecosystem function improved before

earthmoving at the expanded Wherry Dune area, and subsequent substrate

preparation and planting. Removal of these trees will also directly provide

suitable habitat for San Francisco lessingia and its associated species of

concern.

Essential areas for tree removal in dune restoration include upper dune slopes

immediately below Lincoln Boulevard, south of the Baker Beach one-way

entrance. Small groves of trees lower in the dune slope, near the Baker Beach

picnic areas, would be less likely to affect wind velocity upslope. Similarly,

removal of trees at the southwest end of Baker Beach would be unlikely to

affect the integrity of restored Lobos-Wherry Dunes. The removal of trees in

the lower slopes and flats immediately behind the beach would be beneficial

(but not essential) to recovery. On dune slopes immediately above Baker

Beach, which are adjacent to abundant relict native dune vegetation, natural

recolonization of native species in tree-cleared dunes should be fostered before

plans for transplanting of nursery-grown native vegetation are developed. 

Temporary dune surface stabilization (such as desiccated, dead beachgrass)

would initially be needed to enable native seedlings to establish after tree

removal. Buffer plantings of native woody dune scrub species, however,

would need to be placed along Lincoln Boulevard to stabilize devegetated

dunes. 

The Presidio Trust’s goal for removal of Wherry Housing (Presidio Trust

2002) is compatible with timely phased restoration and integration of the

Lobos and Wherry Dune sites. Retention of the housing would have prevented
157

recovery of San Francisco lessingia. Phasing demolition to begin in

approximately 10 years in the area outside of Pershing Drive, and to be

completed in about 30 years, will enable the Wherry and Lobos Dunes units to

become fully integrated. In contrast, occupancy of all of the Wherry Housing

for a full 30 years, instead of phasing, would obstruct recovery of San

Francisco lessingia within this recovery unit because it would preclude

integration (migration, gene flow, colonization of newly available patches)

with the core of the unit at Lobos Dunes. Prolonged occupancy of the Wherry

housing would also set up land use conflicts for full dune restoration including

active blowouts in adjacent dunes, by creating wind-sheltered zones in the lee

of housing, and by establishing land uses that would be intolerant of adjacent

mobile sand. Monitoring of sand movement after removal of buildings should

be instituted as part of an adaptive management framework. 

The final restoration designs of the Wherry Dunes and Baker Beach areas

should anticipate and encourage the development of dune blowouts of

appropriately limited size and distribution. Blowouts should be monitored for

early indications of excessive rates of sand transport or excessive spread of

erosional areas because large, highly mobile blowouts would be detrimental for

San Francisco lessingia, road maintenance, and recreational uses of the park. 

Most blowout migration rates should not normally exceed an average of 0.5

meter (1.6 feet) per year for more than three consecutive years, depending on

location and adjacent vegetation and land management. The dunes at this

locality are unlikely to become mobilized on a large scale because dune soils

here are mature and weathered, and are rapidly colonized by vegetation unless

chronic trampling disturbance becomes intensive. Early intervention can

control excessive blowouts by: (i) planting locally appropriate dune vegetation

in upwind source areas within blowouts during the rainy season of average to

above average rainfall years; or (ii) placing coarse organic debris (e.g.,

desiccated beachgrass straw, partly degraded woody brush) in upwind source

areas and downwind crests of blowouts during drought years.

(3) Rob Hill Dune Remnant. The Rob Hill segment of the proposed LobosWherry dunes complex (near Battery McKinnon-Stotsenberg) supports an

important remnant colony of San Francisco lessingia that should be protected,

enhanced, managed, and expanded within the site’s constraints. The site
158

currently is probably too small and isolated to rely on self-sustaining natural

disturbance dynamics. Removal of nonnative trees at the southwest corner of

Rob Hill would improve exposure to drying, erosional winds, and sun, thus

improving habitat conditions and reduce reliance on intensive management. 

This site, however, should remain under intensive vegetation management at

least until vegetation and San Francisco lessingia population trends indicate

less need for it. In the interim, before habitat is expanded, manual weeding

efforts should be increased. The forest adjacent to this site, the Rob Hill

Buffer, will be managed and retained as historic forest but will also be

managed as a buffer for the San Francisco lessingia population. Management

will include suppression of nonnative invasive plants according to the Presidio

Vegetation Management Plan. San Francisco lessingia and associated native

species should be allowed to invade opened sand habitat where trees and ivy

are removed, while nonnative plant invasions should be suppressed in these

disturbed sites. The Rob Hill reserve may be used to supply additional

founders for new San Francisco lessingia populations in the Wherry-LobosBaker dune complex.

(4) Public Health Services Hospital Dune Remnant. The sandy flats northeast

of the Public Health Service Hospital (former Marine Hospital site) near 15th

Avenue (Figure 11) support relict dune scrub and a rare example of coast live

oak on old dunes, as well as substantial but small colonies of San Francisco

lessingia in disturbed sandy areas (former road and borrow pit, and dune scrub

restoration sites). These colonies, and the small Presidio Golf Course colony

nearby on Park Boulevard, are probably associated with the historic records of

San Francisco lessingia near Mountain Lake, and were probably also originally

linked to the remnant colonies west of Battery Caulfield Road and Lobos

Creek. For this reason, these colonies are important to the conservation of the

species, and should be protected, managed, and enlarged by restoration of

degraded areas within their vicinity. A large proportion of the flats at the

northwest end of the site are affected by high groundwater and seeps that have

developed vegetation typical of coastal dune slacks. This area is not San

Francisco lessingia habitat and should not be restored as such. It would be

highly feasible and appropriate to restore this site as coastal dune slack (dune

wetland) to support species of concern and an open northwest wind corridor

and buffer area, which should be incorporated in the reserve. 
159

The slopes above the San Francisco lessingia colonies are dominated by

iceplant. The nonnative tree grove and iceplant understory must be removed to

expose underlying sand and restore wind-fetch. Implementation of dune

restoration in phases, and use of temporary inert stabilizing materials prior to

revegetation, should prevent excessive erosion following tree removal. 

Removal of planted trees along the east side of Battery Caulfield Road may

reestablish low levels of seed dispersal of San Francisco lessingia from the

Lobos Dunes area to the Public Health Services Hospital population, which

would benefit the population. It would also be beneficial to translocate seed of

San Francisco lessingia from the isolated, but adjacent, golf course roadside

cut to restored sand slopes behind the Public Health Services Hospital because

orientation to effective winds and narrow dispersal corridors are unfavorable

for dispersal of seed from relict sources to new habitat areas. These population

patterns are likely the result of past habitat destruction.

Like the Rob Hill Buffer, much of the Public Health Services Hospital Buffer

is not well suited to self-sustaining disturbance dynamics because of its

relatively sheltered physical setting. It will probably require a relatively high

degree of active vegetation management. The sandy south-facing slope behind

the Public Health Services Hospital is covered with a grove of nonnative

Monterey pine and Monterey cypress. The upper slope also includes large

stands of acacia, iceplant, and tea-tree (Leptospermum laevigatum). This area

will be managed as a buffer to the adjacent San Francisco lessingia populations

according to the Presidio Vegetation Management Plan. This management

includes control and suppression of nonnative invasive plant species.

(5) Presidio Golf Course. The Presidio Golf Course population of San

Francisco lessingia occurs along a road cut in dune sand and is relatively small. 

It is now managed by The Presidio Trust in collaboration with the National

Park Service. This colony may be considered to be an isolated part of the

Public Health Services Hospital remnant population. It lacks adjacent

restorable habitat as long as the golf course remains in use. It has little

potential for long-term conservation except with intensive management. The

population should be conserved in the interim as a hedge against unforseen

catastrophic declines in core reserves, but is not a strategic focus for perpetual

maintenance or expansion. Seed from this population should be incrementally
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reintroduced to the Public Health Services Hospital colonies. Once the Public

Health Services Hospital restoration site has established a persistent population

and manageable successional trends in an estimated 10 to 15 years, the

Presidio Golf Course population should require no extraordinary protection,

maintenance, or expansion beyond what Golden Gate National Recreation

Area has done in the past. It should be conserved as long as feasible, however.

(6) Presidio vegetation management planning. Site-specific planning is

needed in the Presidio Recovery Unit as described in the Presidio Vegetation

Management Plan to develop the measures needed to reach recovery while

maintaining as much as possible the character of the historic forest and meet

other objectives of the Presidio Vegetation Management Plan. The planning

should establish appropriate specific long-term vegetation objectives for the

dune scrub communities supporting San Francisco lessingia. Large areas of

dune scrub should include a significant proportion of vegetation gaps (bare

soil) and patches of sparse vegetation dominated by native annual plant

species. In the Wherry-Lobos Dune complex, the site-specific plans should

stress the dynamic succession of dune blowouts in various stages (including

bare eroding sand). The plans should avoid excessively protecting the area

from trampling and erosion because too much protection can sometimes

promote vegetation succession to closed perennial vegetative cover, especially

in wind-sheltered locations. The plans should include provisions for

experimental use of managed pedestrian trampling and rotational trail closures

and openings as tools to establish vegetation gaps or blowouts. The plans

should prescribe intervention if woody or perennial species become

excessively or uniformly dominant over large areas, resulting in high percent

vegetation canopy cover with few, small gaps. Conversely, if trampling

pressures or dune mobility become locally excessive, affected areas should be

(temporarily) closed to public access, allowing populations of annual plants to

regenerate.

The site-specific plans should prohibit vegetational regeneration of nonnative

trees within restored or managed dune scrub areas dedicated to conservation of

San Francisco lessingia. The plans should also require adaptive management

approaches to invasive weeds, based on frequent surveys to detect incipient

colonies or rapid expansions, eradication of incipient colonies, and confining
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invasion fronts from core weed populations. Weed control should emphasize

(in addition to nonnative trees) iceplants (Carpobrotus edulis and hybrids,

Conicosia pugioniformis), annual bromegrasses (Bromus spp.), oats (Avena

spp.), foxtail (Vulpia bromoides), and, for some localities, Bermuda-sorrel

(Oxalis pes-capreae).

(7) Maintenance and monitoring. All Presidio Dune restoration sites and San

Francisco lessingia subpopulations should be monitored annually to determine

trends of: (i) vegetation succession; (ii) reinvasion by nonnative vegetation;

and (iii) changes in distribution and abundance of San Francisco lessingia. 

Monitoring should include sampling using relatively fine-scale measurements

of percent cover of bare sand, leaf litter, and live vegetation, as well as erosion

and accretion around San Francisco lessingia colonies and adjacent unoccupied

sites. Monitoring should include periodic low-elevation color infrared aerial

photography of the Presidio Recovery Unit to enable accurate vegetation maps

to be constructed. Monitoring of restored dune areas within complexes should

include some detailed subsampling of San Francisco lessingia population

dynamics using demographic methods. Monitoring of San Francisco lessingia

should compare its population dynamics within blowouts, at blowout edges,

and in higher density dune scrub and grassland vegetation. Intensive

population data collection, however, should not be applied indiscriminately for

monitoring large reserves when more efficient sampling methods would suffice

for management.

We, the U.S. Fish and Wildlife Service, recommend that at least one full-time

vegetation technician qualified to maintain restored dune areas should be

retained cooperatively by the Golden Gate National Recreation Area and

Presidio Trust to integrate long-term monitoring, management, and

maintenance activities in the restored dune system supporting San Francisco

lessingia. The site-specific plans developed to implement the Presidio

Vegetation Management Plan should be updated to adapt to the management

needs of this unit. We recommend that a dedicated management fund for

restored areas supporting San Francisco lessingia should be established within

the Presidio Trust and Golden Gate National Recreation Area Park budgets to

ensure that habitat gains from reintroduction in the Presidio are not

subsequently lost during periods of budget reduction.
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b. Southern Recovery Unit.

i. Daly City Reserve. The Hillside Park sand slope (Daly City) population

occurs partly on land owned by the City of Daly City and partly on privatelyowned lots that have remained undeveloped on this steep, erosion-prone sandy

slope. Although dense stands of San Francisco lessingia are confined to the

vicinity of a pipeline and disturbed areas around foot-trails and open sand, San

Francisco lessingia has at least recently (1998) spread over most of the area to

the west and east. Habitat suitability for San Francisco lessingia here is

probably unstable in the long term, and will require management of vegetation

in order for San Francisco lessingia to persist there.

At the Hillside Park sand slope, the current property owners have neither the

mandate nor resources to conserve San Francisco lessingia effectively. 

Therefore, recovery will entail either: (a) acquisition and transfer of this site

(including restorable habitat and buffer areas) to a conservation agency or

qualified organization dedicated to conservation of San Francisco lessingia and

its native plant community; or (b) establishment of enforceable agreements

(possibly easements) to manage the site cooperatively as a reserve for San

Francisco lessingia and associated vegetation, coupled with an endowment for

management and maintenance to ensure the resources needed for management. 

Of the two options, acquisition is preferable because it reduces complexity and

uncertainty of management. Acquisition should occur through steps including

public outreach to neighboring residents, identification of willing sellers, and

identification of potential long-term managers. 

The areas adjoining the San Francisco lessingia population at Hillside Park

support restorable sandy habitats (under mixed native and nonnative

vegetation) and plant associations typical of coastal dune scrub. For example,

suitable sparsely vegetated and disturbed sand slopes occur next to a public

school at the east end of the hill. These slopes afford potential for population

expansion, conservation, public stewardship, and scientific education. These

areas should be integrated into a comprehensive preserve with different

management units. Management units would include: (a) perimeter buffer

zones of dune scrub; (b) core Lessingia population area (a priority area for

control of invasive nonnative plants); (c) stable dune restoration sites
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(currently dominated by nonnatives, subject to future lessingia expansion or

reintroduction); and (d) dune scrub preserves.

A management and restoration plan for the Hillside Park site should be

developed in cooperation with local government and neighboring residents

with interest in public access to the site and its educational values. The aims of

the management and restoration plan should include: (a) promoting and

maintaining sparse, relatively open dune scrub vegetation favoring annual gapcolonizing species, in particular San Francisco lessingia in areas it currently

occupies; (b) restoring open dune vegetation in adjoining areas unoccupied by

San Francisco lessingia that are dominated by nonnative vegetation; (c)

facilitating dispersal and colonization of San Francisco lessingia in restored

areas; (d) establishing managed vegetation buffers to slow reinvasion by

invasive nonnative species that degrade San Francisco lessingia habitat; and (e)

ensuring that all parts of the management unit are efficiently designed and

functionally integrated. Management plans should incorporate recovery

criteria for this reserve.

Management and restoration plans for the consolidated Hillside Park dune

scrub vegetation preserve must include: (a) provisions for ongoing suppression

of invasive vegetation within areas currently occupied by San Francisco

lessingia, utilizing manual removal techniques or grass-specific herbicides as

appropriate; (b) geotechnical assessment of slope stabilization needs; (c)

assessment of existing infrastructure (e.g., buried utility lines) constraints, and

provisions for any re-routing or replacement of infrastructure to make it

compatible with long-term vegetation management; and (d) gradual, phased

removal and eradication of iceplant and annual alien grasses, and planting of

buffer vegetation comprising native dune scrub species along the edge of the

site to retard seed dispersal of annual grasses back onto managed dune

vegetation. 

Regulated public access to the Daly City Reserve would enable natural history

interpretation and provide educational opportunities. Public access designs

and interpretative signs for dune scrub habitats should be modifications of

those at the Presidio’s Lobos Dunes and Crissy Field dune/salt marsh
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restoration projects, which were designed by the Golden Gate National Parks

Conservancy and the Golden Gate National Recreation Area. 

Within restored areas, adaptive management plans should prescribe

experiments with seasonal closures and openings of pedestrian access to rotate

trampling disturbances in sites dominated by dune annuals. This “managed

trampling” should be scheduled in the fall, after seed dispersal is relatively

complete, but before germination begins. The site and site manager should be

endowed with a maintenance capital fund to ensure that sufficient funds will be

generated annually to weed and otherwise manage the site.

ii. Fort Funston Reserve. In the southwest corner of San Francisco County at

south end of the Golden Gate National Recreation Area are Fort Funston and

an area of relictual dunes just to the north. The Fort Funston Dunes are owned

and managed by the Golden Gate National Recreation Area. They represent

the only large remnant dune habitat that occurs directly within the historic

southern portion of San Francisco lessingia’s range. The vegetation here

includes remnant dune scrub, restored early succession dune scrub, groves of

nonnative vegetation (Monterey cypress and pine, acacia, and plume acacia

[Albizia lophantha], and extensive areas dominated by iceplant) and bare sand. 

The Fort Funston dunes are relatively heavily trampled in many areas, and are

a highly popular dog exercise area. At least 40 hectares (100 acres) of existing

suitable habitat and restorable habitat for colonization by San Francisco

lessingia are estimated to exist at Fort Funston, giving the area the potential for

a major population. Because intensity of recreational use tends to diminish

with distance from parking areas and paved paths, the northern end of the

Funston dunes (approximately 26 hectares [65 acres]) should be given highest

restoration priority. Major efforts here should be large scale removal of

nonnative invasive vegetation, particularly weedy annual grasses and iceplant,

and reintroduction of San Francisco lessingia and associated dune scrub

species, including native grasses and shrubs. Toward the south end of the

dunes within the Golden Gate National Recreation Area, less intensive

management and restoration effort would be appropriate. The semi-disturbed

nature of the southern area and the ruderal (weedy) tendencies of San

Francisco lessingia probably make reintroduction of this species compatible

with the park’s mixed recreational and conservation land uses, so
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reintroduction should not require significant additional restrictions on existing

recreational uses. San Francisco lessingia should be reintroduced to the Fort

Funston Reserve in phases, using founder colonies in temporary small

exclosures (fenced areas designed to keep out people or animals). Areas

subject to reintroduction in this area should be at least 4 hectares (10 acres). 

Within the Fort Funston Reserve, buffers around historic structures should be

managed to avoid detrimental effects to the adjacent dune scrub habitat. We

recommend that studies be done to determine recreational use management

consistent with maintenance of dune scrub habitat and sustainable populations

of San Francisco lessingia.

To the south of Fort Funston and the Golden Gate National Recreation Area is

an open space area in private ownership. We believe this site may include

areas of suitable topography where dune scrub habitat could be restored. If

funding is available and the landowners are interested, weed control and

restoration in the area could contribute to the recovery of San Francisco

lessingia. Such measures might also provide some small increases to the areas

available to the recreationists who enjoy the dune scrub areas in the Fort

Funston dunes. Increased recreational use may benefit any restored dune scrub

areas in suitable condition for growth of San Francisco lessingia by

maintaining disturbance.

The first phase of restoration and reintroduction would include cultivation of

lessingia transplants at the Fort Funston nursery, and direct sowing into

selected relatively bare sand areas already protected from heavy trampling

(such as existing dune scrub restoration sites). The founder populations should

be derived from the Hillside Park (Daly City) population, which is genetically

distinct from the Presidio population (Staci Markos, Jepson Herbarium, U.C.

Berkeley, in litt., 2002), is likely related to the population formerly found at

Fort Funston, is susceptible to extirpation due to its small size, is

geographically close to Fort Funston, and occurs in habitat that is similar in

sand/soil characteristics. Seed should be sampled extensively throughout the

Hillside Park parent population to allow for high initial genetic diversity on

which subsequent natural selection or genetic drift may act after reintroduction.

Transplants should be made during rainy weeks in winter. Founder

populations should comprise at least 200 transplants and several thousand
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direct-sown seed, distributed over areas ranging in size from about 0.05 to 0.1

hectare (0.12 to 0.25 acre). Transplanting should be postponed during winters

with low, infrequent rainfall. Growth rates of transplants should be augmented

by addition of soil nutrients (particularly nitrates) to containers shortly before

transplanting.

The second phase would begin once founder populations are established and

expanding spontaneously. It would involve removal of iceplant mats (from

areas with minimal native vegetation), to avoid competition with dune scrub

vegetation. Iceplant removal should also include the full removal of the semidecomposed organic layer beneath the live tissue in order to establish good

sand conditions. Iceplant-cleared plots approximately 0.4 to 1.2 hectares (1 to

3 acres) in size should be replanted patchily with native dune scrub species

propagated from local stock. Founder populations of San Francisco lessingia

derived from Phase 1 populations should be similarly established in these

areas. Once the native species are established and spreading after about 3 to 5

years (depending on climate-driven variables), restored dune scrub areas with

reintroduced San Francisco lessingia may be re-opened for recreational access,

either on rotational trails or without trail restrictions, as park managers

determine appropriate based on monitoring and preliminary studies. 

Exclosures should be needed only if managers detect widespread severe

trampling-related mortality of San Francisco lessingia. If native scrub species

develop dense and relatively continuous cover, or if iceplant or annual grasses

reinvade restored plots excessively, they should be removed to control

vegetational regeneration and protect the semi-open nature of the vegetation

that is conducive to San Francisco lessingia’s persistence. 

 Vegetation management aims for the Fort Funston Reserve should be based on

the area occupied and the total population size of San Francisco lessingia, both

of which would fluctuate among years. Initial performance over the first 10

years should aim for at least 5 colonies, each occupying a variable area, with

the overall population ranging from approximately 100,000 to 500,000 or more

individuals. The Fort Funston Reserve should be monitored and maintained

according to the general prescriptions for the Presidio Unit. A dedicated

management fund for restored areas supporting San Francisco lessingia should

be established within the Golden Gate National Recreation Area Park budget to
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ensure that habitat gains from reintroduction at Fort Funston are not

subsequently lost.

c. Satellite Recovery Unit. It is possible that catastrophic events (e.g., fire, disease,

insect outbreaks, etc.) at the Presidio and Fort Funston could cause extinction of the

species. To reduce this risk, a series of smaller separate satellite populations should be

introduced to independent reserves on restored remnant dune habitats in Sunset

Heights and Sutro Heights (near the Cliff House, sand slopes above Balboa Street and

48th Avenue). These sites contain plant species regularly associated with San

Francisco lessingia, as well as large stands dominated by nonnative vegetation. These

scattered sites are within the ecological and geographic range of San Francisco

lessingia, but are not specific historic collection localities. Reintroduction of San

Francisco lessingia to restored areas at these locations would help compensate for the

loss of the Lone Mountain population of San Francisco lessingia (all contain hills of

Franciscan bedrock, with climbing dune veneers), and would further spread the risk of

extinction among more independent populations. 

i. Sutro Heights Sub-Unit. Portions of the Sutro Heights site (lower slope) are

privately owned, but much of the area is owned by the City of San Francisco

(Department of Parks and Recreation). The Sutro Heights dune remnant site is

near the Presidio population, but is similar in soil characteristics (Colma

Formation and dune veneer) to the Merced/Oceanside dunes locality where San

Francisco lessingia occurred historically. Establishing a new population here

would provide geographic and ecological variation for the species that is not

provided in the main recovery units. Conditions at this site are comparable to

those at the historic (now extirpated) colony at the west end of Baker Beach. If

it becomes well established at the Sutro Heights location, San Francisco

lessingia might spread spontaneously to some stabilized portions of the

engineered dune ridge along Ocean Beach and the Great Highway. It would

not establish in areas of mobile sand. The original relict population of San

Francisco lessingia near Lobos Creek occurred near a derelict roadside, and

other Lessingia species are known to behave as weedy invaders of sandy

roadsides (Spence 1964). Any adventive colonies spreading from the

reintroduced population along Ocean Beach/Great Highway should be treated

as dynamic, transient populations; normal maintenance and recreational
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activities should not be prohibited in attempts stabilize individual colonies

here.

Recovery actions for Sutro Heights dunes include removal of nonnative woody

vegetation (acacia, plume acacia, Monterey cypress) and iceplant, and

stabilization with native dune grassland species. Interim physical stabilization,

long-term stabilization by vegetation, and buffers are necessary because of

adjacent residential land uses. Site preparation should include inventory and

salvage of existing and spontaneously recruited (seed bank) native plant

species that may emerge during nonnative vegetation removal. Preparation

must also include eradication of invasive nonnative species prior to

reintroduction of San Francisco lessingia. San Francisco lessingia from the

Presidio population should be used as founders of the new colonies at Sutro

Heights because they are close geographically, and because the Presidio plants

were historically collected from equivalent ecological conditions (sprayexposed Colma Formation sandy bluffs; Table 1). Restoration and

reintroduction methods should approximately follow those of Fort Funston.

ii. Sunset Heights Sub-Unit. Sunset Heights sites also have some potential for

joint reserves for experimentally established populations of Raven’s manzanita

where Franciscan bedrock outcrops (including greenstone) occur. The Sunset

Heights dune remnants occur at Grandview Park, a west-facing slope of Sunset

Heights Park, an area of mixed outcrops and dunes above 15th Avenue and

Ortega Street, and “Hawk Hill” (near 14th Avenue and Santiago Street, above

the Herbert Hoover Jr. High School). The City of San Francisco owns

Grandview Park and Sunset Heights Park, and has recently acquired the “Hawk

Hill” site. Hawk Hill also supports many dune plant species of concern and

regional conservation significance.

The first recovery actions for the Sunset Heights Sub-Unit must be to secure

the sites for conservation of dune vegetation and reintroduction of San

Francisco lessingia. Cooperative agreements should be established with the

City of San Francisco Department of Parks and Recreation to manage invasive

nonnative vegetation (eradicate iceplant and tall veldtgrass [Ehrharta erecta]

in particular), and to cut back nonnative woody vegetation encroaching the

margins of the existing dune remnant at Sunset Heights Park. A vegetation
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management fund should be established for the maintenance removal of

nonnative invasive vegetation at these sites. 

It is not clear whether seed to found the new Sunset Heights population should

come from the Hillside Park (Daly City) population, or from the Presidio

population. Sunset Heights is slightly closer to the historic Lake

Merced/Oceanview localities, which may have given rise to the Daly City

population, but its dune soils and local climate are more similar to those of the

Presidio. Identifying an appropriate seed source will require preparation of a

genetics management plan following the guidelines of the Center for Plant

Conservation, as required by our policy. Further investigation of the genetics

of the species and the two strains may be needed. We will adapt the

management of San Francisco lessingia as we learn more from genetics

management planning, investigations, and experience in the field.

Given the outcrossing nature of San Francisco lessingia reproduction and the

genetic distinctness between the Presidio and Daly City strains, establishing

populations of plants with a mixture of parentage from the two types may

either be beneficial (outcrossing plant species often gain vigor when dissimilar

strains are allowed to breed) or detrimental (crossing different strains can

depress vigor if they have become too distinct). Possible options to identify an

appropriate seed source for Sunset Heights through the genetics management

plan include: a) Conducting preliminary experimental transplants from both

strains and monitoring them on site to determine whether adaptively significant

differences can be detected between source populations; b) testing

within-species "hybrids" between parents from Daly City and Presidio

populations to compare growth and reproductive output of the hybrids with

parent populations; and c) greenhouse testing of Presidio and Daly City

crosses (isolated from propagation connected with re-establishment) to screen

for problems due to poor hybrid viability from such crosses. The seed source

that proves best adapted under such tests should be used for reestablishment. 

However, if shortages of funding would cause a long delay in such

investigations, the genetics management plan should identify a preferred strain

to be used and reestablishment efforts at Sunset Heights should proceed. If

reestablishment efforts fail, the other strain or crosses between the strains can

be tried. These decisions should follow an adaptive management strategy laid
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out in the genetics management plan for San Francisco lessingia. Preparation

of this plan should therefore proceed as a high priority due to the need to be

ready to proceed with reestablishment seeding as soon as each patch of habitat

is restored.

Management plans for vegetation at satellite reserves should be prepared and

implemented with public notice and participation. The Hawk Hill site should

be managed by the City of San Francisco (Parks and Recreation), or by a

nonprofit conservation trust or government resource agency. Privately owned

portions of the Sutro Heights sand slopes should be acquired and transferred

either to the Golden Gate National Recreation Area, San Francisco Department

of Parks and Recreation, or a qualified nonprofit conservation trust. This site

also should be endowed with funds for maintenance in perpetuity. The sites

should be monitored following the general prescriptions for the Presidio Unit. 

The satellite populations should be treated initially as though experimental

populations. 

2. Recovery Strategy for Raven’s Manzanita 

The recovery of Raven’s manzanita, like that of San Francisco lessingia, is based on

dual conservative and experimental approaches, emphasizing both protection of

existing populations in enhanced existing habitat, and experimental establishment of

new populations in restored habitat. The Presidio Unit’s recovery actions are

primarily conservative: they stress the survival of the remnant natural clone and its

daughter clones, with vegetation management to suppress excessive competition from

invasive nonnative plants. The San Francisco Unit’s recovery actions are primarily

experimental, though probably feasible, based on horticultural experience with native

San Franciscan manzanitas. These actions consist of: (i) selecting undeveloped

(usually undevelopable) rock outcrop sites (mostly serpentine) in the city that support

either sparsely vegetated, bare, or predominantly weedy vegetation; (ii) preparing the

sites for reintroduction of native manzanitas (including Franciscan manzanita) and

native associates; and (iii) establishing populations of new genetic individuals of

Raven’s manzanita and associated species of rock outcrops (particularly serpentine). 

Recovery actions at interior San Francisco sites also include measures to minimize

pollen flow from planted ornamental manzanitas. These replicated mixed populations

of native manzanita would be actively maintained during establishment periods, then
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allowed to grow (or fail) independently, with low-level maintenance. They would be

monitored for many years to track growth and (expected) sexually reproduced

generations. 

This recovery approach has two important milestones: (i) establishment of additional

replicate clones of the remaining wild Presidio plant at many independent sites, not

just clustered around the parent plant, to avoid extinction from localized events; and

(ii) establishment of multiple generations of genetically-variable individuals in

sexually reproducing populations. The first steps toward spreading the risk of

extinction among multiple cloned populations of the Presidio plant have been taken by

the Golden Gate National Recreation Area and the U.S. Army, which have already

planted 50 daughter clones near the mother clone and at 3 other Presidio locations

(Golden Gate National Recreation Area unpublished data). However, most of these

are subject to risks of spatially dependent mortality such as disease, fire, vandalism,

trampling, and insect damage. This initial effort needs to be extended by more

widespread establishment of additional daughter clones to variable serpentine outcrops

of the Presidio.

New genetically variable populations should be established to approximate the

unknown number of extirpated populations of Raven’s manzanita from interior San

Francisco bedrock outcrops (at least four have been documented). The new

populations should be in a variety of microclimates, generally warmer and drier than

the Presidio. Because of the scarcity of suitable reintroduction sites, removal of

nonnative trees will probably be necessary for reintroduction experiments. New

populations of Raven’s manzanita should be established by sexual reproduction (seed)

in order to produce variable new genetic individuals (as opposed to clones from

cuttings). The first priority for recovery should be to establish viable inbred lines of

Raven’s manzanita, if possible (D. Elam, pers. comm. 1999), through artificial selfpollination in controlled conditions. The ratio of viable seed per self-pollinated

flowers, the germination and survival rates of any inbred seedlings, and the variability

of any surviving inbred plants (genetic, morphological, and growth characteristics in

variable substrates and microclimates) should be investigated. Any new inbred

genetic individuals should be carefully labeled, catalogued, and kept in cultivation. 

Individuals determined to be suitable for reintroduction should be propagated clonally.
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If it proves infeasible to produce adequately fit offspring that are genetically variable

through self-pollination of the single surviving Presidio clone, it will be necessary to

either abandon the objective of achieving sexually reproducing populations of Raven’s

manzanita, or try alternative breeding methods to generate sexually reproducing

populations with increased genetic variability. Breeding strategies based on

introgression, or recurrent selection of introgressed hybrids (see Recovery Strategy,

page 75) could be attempted with closely related taxa that are ecologically and

morphologically similar to Raven’s manzanita, emphasizing individual parent plants

that are particularly similar. These strategies should be evaluated as described above

for inbred Raven’s manzanita. Recommendations on any potential breeding programs

involving Raven’s manzanita should be made by a scientific peer review panel with

expertise in manzanita biology, plant conservation genetics, and plant breeding, after

new scientific data on species’ systematics and reproductive biology (described in

recovery criteria) becomes available. The panel should also develop a genetic

management plan for the species. Any experimental backcrossed hybrid derivatives of

Raven’s manzanita produced must be kept carefully isolated from Presidio

populations. Tamalpais manzanita is presumably the most appropriate potential

hybrid parent (based on current understanding of manzanita taxonomy, biogeography,

and ecology) but other manzanita species should be evaluated as well, based on prior

molecular genetic, ecological, and morphological comparative studies. 

Other recovery actions include investigation of the levels of genetic variation in

Raven’s manzanita compared with small populations of rare wild relative manzanitas;

evaluation of new genetic individuals’ ecological, morphological, and physiological

variation; investigation of taxonomic relationships between this and related manzanita

species; and horticultural research on propagation, transplanting, and establishment. 

Public education and outreach are also essential actions that must be coordinated with

implementation of recovery actions involving habitat restoration and reintroduction.

a. Development of a New “Pure” Inbred Founder Population of Raven’s manzanita.

Since no natural seedling establishment of Raven’s manzanita has ever been observed

since it was rediscovered by Peter Raven in the 1950's, artificial seed propagation is

the only potentially practicable way to obtain new genetic individuals to add to the

population. Seedling survival rather than seed germination itself has limited

production in past efforts at growing seed from open-pollinated Raven’s manzanita (H.

Forbes, pers. comm. 1999), so emphasis should be placed on: (i) ensuring that seed is
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produced exclusively from controlled artificial self-pollination; (ii) providing

intensive, high quality seedling aftercare of any inbred seedlings; and (iii) promoting

rapid growth and early cloning of new juvenile plants. Experimental treatments that

shorten the juvenile phase and promote precocious sexual reproduction would benefit

development of inbred lines. Annual intensive controlled pollination and seed

production, seed collection and propagation of cultivated clones will be needed to

augment seedling production. 

Any new genotypes of inbred Raven’s manzanita should be clonally replicated at

multiple botanical gardens and the Golden Gate National Recreation Area’s native

plant nurseries, with mandatory permanent labeling and identification of clones and

their pedigrees. New individuals should be maintained in permanent cultivated

populations. If they are determined to be fit for use as stock for outplanting in new

populations in San Francisco outside of the Presidio, replicated clonal stock

populations should be maintained in cultivation. Golden Gate National Recreation

Area nurseries at Fort Cronkhite and the Presidio would be well-suited facilities for

the horticultural propagation and any breeding work. In addition, replicates of

existing and future clones of inbred and cloned “pure” Raven’s manzanita should be

maintained in perpetuity at multiple botanical gardens in California, in relative

isolation from other manzanita species. Seed of any inbred Raven’s manzanita should

be stored at local (Golden Gate National Recreation Area, University of California

Botanical Garden, Berkeley) and other (Rancho Santa Ana) seed banks, following

guidelines of the Center for Plant Conservation (1991). Cloned new genotypes should

be tested for performance (growth, reproduction) in propagation, and in new interior

San Francisco reintroduction sites.

b. Contingency Measures for Producing New Individuals: Introgressive Breeding of

Raven’s Manzanita. It is possible that self-pollination of clones of the single

remaining individual may produce viable seed, and enough seedlings with adequate

variability in ecologically important traits (e.g., growth habit, growth rate, soil

chemistry tolerance, disease resistance) for natural selection to act upon. This

outcome may be possible if the single clone is not a fully obligate outcrosser, and if it

has a relatively high amount of latent genetic variation (many different forms of genes,

or alleles, and multiple alleles for various genetic traits) as was the case for the singlefounder, endangered showy indian-clover (Trifolium amoenum) (Knapp and Connors

1999). If, however, selfing produces no seed, or plants with inadequate fitness or poor
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adaptability to environments of reintroduction sites (e.g., high post-establishment

mortality, weak competitive ability, unusually slow growth, disease susceptibility,

nutrient deficiency symptoms in local serpentine soil), an alternative source of genetic

variation may be developed by an introgressive breeding strategy (see Recovery

Strategy, page 75). Introgressive breeding (Falk 1992) would be based on creation of

hybrids with similar, closely related manzanitas, selection of hybrids, and

backcrossing (crossing hybrids with one parent species) on to Raven’s manzanita over

multiple generations. Backcrossed individuals that are essentially indistinguishable

from Raven’s manzanita species (as in examples of natural introgression among other

manzanitas; Ellstrand et al. 1987), but exhibit a wider range of variability, would be

selected repeatedly. This variation on a “captive breeding” strategy for plants is a

tactic of last resort, justified only when populations are so reduced that reproduction of

genetically viable generations is practically infeasible (Rieseberg 1991, Falk 1992). 

Introgression among manzanita species occurs in nature, and in some cases results in

individuals that are essentially indistinguishable from parent taxa, even in specialized

serpentine species. Gottlieb (1968), examining populations of serpentine-endemic

whiteleaf manzanita (Arctostaphylos viscida), found only a few of its hybrids on

serpentine soil, and noted that these presumed introgressants closely resembled A.

viscida. Ellstrand et al. (1987), studying natural hybridization in A. viscida ssp.

mariposa and A. patula, found that most introgressants were physiologically similar or

indistinguishable from parent species. They also found that true hybrids (firstgeneration species crosses) and segregants (progeny of hybrids breeding among

themselves) were rare. Kruckeberg (1977) observed that the frequent generation of

hybrid Arctostaphylos ×media (from A. columbiana and A. uva-ursi) did not appear to

cause genetic contamination of the parent species by gene flow through introgressants. 

In the absence of experimental data from Raven’s manzanita, it is unknown whether

these natural analogues are relevant to the specific case of Raven’s manzanita. The

decision to introduce artificially bred introgressant forms of Raven’s manzanita to

isolated, newly restored urban reserves must involve scientific peer review by

botanists with expertise in manzanita systematics and ecology, and in population

genetics of rare plants, based on research outlined above.

A comparable “single founder” challenge to endangered plant species recovery has

been recently investigated in the showy Indian clover (Trifolium amoenum), a species

federally listed as endangered, and a “presumed extinct” species (Isely 1993) that was
175

believed to have been reduced to a single known fertile individual when it was

rediscovered at an inland location in Sonoma County, California. A second wild

coastal population of 20 plants was subsequently discovered. In this case, a surprising

amount of genetic variation (as polymorphism of alternative genes [alleles] for forms

of proteins called allozymes) was detected within the lone survivor of the inland

population (Knapp and Connors 1999). Over 50,000 viable seeds were derived

quickly from self-pollination of the single known surviving plant, and seedlings

thrived in cultivation. In this case, strong ecological and morphological contrasts

between the two known populations suggested that benefits of increasing genetic

diversity by simple hybridization of distinct inland and coastal ecotypes (ecological

races) would be outweighed by the risks of producing maladapted hybrid offspring,

and a decline in fitness (outbreeding depression). This ecological adaptation issue

may be moot for Raven’s manzanita and Tamalpais manzanita, both of which inhabit

serpentine outcrops near the coast around the Golden Gate. The systematic

relationship between these two closely related manzanitas, however, is an open

question (Markos et al. 1999). Additional research will be necessary to determine

whether Raven’s and Tamalpais manzanita constitute one species with distinct

populations, subspecies within a species (as recently interpreted; Wells 1993), or

different species altogether.

The use of hybridization to any degree in the genetic management of an endangered

plant species (even a sole surviving individual) is likely to be controversial since there

are no actual precedents, only proposals. Knapp and Connors (1999) suggested that a

breeding program of controlled experimental introgression that entails repeatedly

crossing a series of hybrids (distinct populations within the species) back on to one

parent population might provide a means of balancing the needs for genetic variation

and adaptive integrity in showy Indian clover. Rieseberg (1991) suggested that

hybridization might be useful in captive breeding programs as a “last-ditch” effort to

conserve endangered species. Barrett and Kohn (1991) recommended experimental

approaches to reintroduction of rare plants that involved both “pure samples” from

existing populations as well as “composite [within species] mixtures with greater

genetic variation,” noting the uncertainty of theoretical predictions regarding genetic

management of “pure” or mixed founders of reintroduced populations. Reintroducing

experimental populations of both “pure” and “mixed” genetic individuals at isolated

locations would enable scientific testing of how well the compromise strategy works

(Knapp and Connors 1999, Guerrant 1996). The appropriateness of the use of
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hybridization in genetic management of Raven’s manzanita may depend, to a

significant degree, on how closely related it is to manzanitas considered as candidates

for hybridization.

Because of the natural ecological and geographic isolation of serpentine outcrops in

interior San Francisco, and lack of natural seedling establishment of Raven’s

manzanita after five decades of observation, it is unlikely that experimental

introgressive breeding, if implemented, would pose genetic risks to the natural coastal

Presidio plant. This assessment is also suggested by the integrity of parent manzanita

species immediately outside the edges of their hybrid zones (Gottlieb 1968,

Kruckeberg 1977, Ellstrand et al. 1987). Any introgressive hybrids of Raven’s

manzanita must be carefully maintained and labeled (with parentage) at Serviceapproved botanical gardens or nurseries dedicated to either research or native plant

restoration. Seed of any introgressive hybrids should be stored at local (Golden Gate

National Recreation Area, University of California Botanical Garden, Berkeley) and

other (Rancho Santa Ana) seed banks, following guidelines of the Center for Plant

Conservation (1991).

c. Site Selection and Acquisition for New Populations in San Francisco. Suitable

candidate sites should be assessed for pilot reintroduction projects. Sites should be

selected based on information obtained by completing detailed field surveys, and

applying the following criteria (in order of priority): (i) presence of serpentine

bedrock outcrops within open space areas that have potential for vegetation

management or restoration, (ii) presence of greenstone (basalts or similar volcanic

rocks with mafic chemistry) outcrops within open space areas, and (iii) presence of

other Franciscan rock outcrops with thin or minimal soil development. Suitable sites

would include steep slopes. Additional factors that may affect selection of candidate

sites include feasibility of nonnative plant removal and subsequent control and

management; minimization of local land use conflicts or local opposition to removal

of nonnative vegetation (especially trees); and local support for vegetation

management. 

Likely areas for candidate sites within the Fort Point/Hunters Point serpentine belt

include Bayview Hills, Potrero Hills, the U.S. Mint at Duboce Street, slopes behind

Crissy Field (Presidio), and slopes between Fort Point and Baker Beach (Fort Scott)

(Figure 2 and Figure 3). Sites would likely include road cuts or old excavations. 
177

Greenstone sites are likely at Mount Davidson Park, and Mount Sutro. Potrero Hills is

a particularly suitable potential reintroduction area. Although it is not a specific

historic locality of manzanitas, it contains some conserved open-space serpentine

outcrop sites within the same serpentine belt as historic populations (Figure 3). Other

sites with likely potential bedrock outcrops suitable for experimental reintroduction of

native manzanitas include mixed Franciscan rock outcrops Corona Heights, Twin

Peaks, Sunset Heights, McLaren Park, and Point Lobos. 

Following site evaluation and selection, sites should be acquired so that habitat

restoration and reintroduction of native manzanitas and their associated rock outcrop

flora can be implemented. Management plans must be prepared for each reserve. 

Maintenance of each reserve should be ensured by selecting an appropriate manager,

initiating local stewardship programs, and endowing the site with a maintenance and

monitoring fund.

d. Control of Cultivated Manzanita Pollen Flow and Spontaneous “Nonnative”

Hybrids. Full recovery of Raven’s manzanita (and reestablishment of Franciscan

manzanita) would require local removal of nonnative manzanitas within likely

pollinator flight distances of transplanted Raven’s manzanita populations to prevent

their reproduction from being “swamped” by pollen from abundant planted

manzanitas, resulting in the Raven’s manzanitas producing predominantly hybrid seed

(Levin et al. 1996). Pollinator distances for significant gene flow among manzanita

populations would be difficult to estimate, and would be geographically specific. 

Scientific estimates should be obtained by site-specific research. Alternatively, a ban

on planting other manzanita species in the western quarter of the Presidio may be used

as a practical surrogate for unavailable scientific data, based on best professional

judgement. Currently, all manzanitas are prohibited from landscape planting on the

Presidio, in order to preserve the integrity of the Raven’s manzanita (Brian O’Neill,

National Park Service, in litt. 2002). Reducing potential “pollen swamping” near

reintroduction sites should be achieved by public education, outreach, and

cooperation, with the assistance of local conservation organizations. In contrast, some

or most transplanted populations of Raven’s manzanita outside the Presidio should be

interplanted with Franciscan manzanita, its historic associated species. Newly

established populations of Raven’s manzanita should be monitored closely for

reproductive output (seed production) and success (seedling establishment) over a

period of decades (multiple generations).
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e. Identification and Permanent Protection of Existing and Potential Serpentine

Habitat in the Presidio. In order to conserve these scarce serpentine outcrop resources

for future potential establishment of native manzanita populations, surface exposures

of serpentine rocks and soils in the Presidio should be: (i) surveyed; (ii) assigned

reasonable buffers in consultation with us under the Endangered Species Act; and (iii)

automatically transferred to the Golden Gate National Recreation Area, if not retained

by the Presidio Trust in the future. Both the Trust and Golden Gate National

Recreation Area should maximize opportunities to conduct habitat restoration and

reintroduction of Raven’s manzanita to these sites. Areas within the Presidio bearing

outcrops or near-outcrops of serpentine bedrock should not be transferred to the U.S.

General Services Administration for disposal. At the present time, legislation

authorizing the Presidio Trust does not preclude such transfers.

f. Public Outreach and Education. Public outreach and education are needed for the

long-term recovery of Raven’s manzanita, primarily to develop support and

cooperation for restoration and reintroduction actions, and to avoid opposition to the

changes in the urban landscape that are necessarily part of restoration projects, such as

removal of nonnative trees. Both tree removal, and reintroduction of species that are

federally listed as endangered, may encounter local opposition, which means that

collaborative outreach and education programs are essential prior to initiation of

recovery projects. These programs can utilize local schools, nonprofit conservation

and horticultural organizations, and should include a program to promote appropriate

ornamental horticultural use of native manzanitas (other than federally or State listed

taxa). Local broadcast and print media can explain the need and purpose of restoration

actions, and might promote the esthetic benefits of improved scenic views following

removal of nonnative trees. Local stewardship programs, many of which exist even

now in potential reintroduction sites, can provide highly valuable long-term

maintenance and monitoring through stewardship programs. Authorized public

demonstration gardens can serve as auxiliary botanical gardens for local

neighborhoods, familiarizing local residents with Raven’s manzanita and providing

reassurance that reintroduction will not cause restrictive regulatory burdens or changes

to recreational land uses. Appropriate horticultural use of endangered plant species as

a tool for public outreach in recovery has been proposed for the endangered western

lily (Lilium occidentale) (U.S. Fish and Wildlife Service 1998).
179

g. Additional Research. Long-term research is particularly needed to determine

appropriate management of the species. Applied research actions in addition to those

identified for breeding and reintroduction (see §IV.B.2.b on pages 173 to 176) should

include ecophysiological studies to evaluate basic environmental growth responses

(e.g. growth under variable soil/bedrock types, mycorrhizal function, competition,

slope, aspect) of the species. Investigation of the reproductive ecology, particularly

seed germination, seedling establishment, seed set, seed dispersal, and pollination, are

needed to predict and manage conditions necessary for spontaneous seedling

recruitment. Practical research needs also include locally appropriate methods of

controlling invasive nonnative vegetation.

3. Conservation Recommendations

The following conservation recommendations are directed at species of concern and

conservation significance. These species are the vanishing components of the local

dune remnants and thinly vegetated bedrock outcrops of San Francisco that were

discussed in Chapter III.B. These recommendations should be integrated with the

recovery actions that are aimed at federally listed species, to assure that the recovery

actions result in community-level protection, management, and restoration actions. 

General tasks for species of concern or conservation significance include the

following:

(a) Survey potential restoration sites, including seed bank probes (germination

tests of shallow soil sample cores) for the presence of target species. Survey

appropriate habitat for target species within San Francisco. Surveys should be

conducted over multiple years, including drought and high rainfall years. If

surveys are negative over several years, or if remnant populations are small

and at least regionally rare, expand surveys to nearest suitable habitat within

the species’ geographic range.

(b) If lands supporting viable populations of species of concern are

unprotected or adversely managed or neglected, acquire or protect parcels as

reserves. 
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(c) Enhance habitats supporting target species where they have been degraded

by influences such as invasion by nonnative plants, excessive trampling or offroad vehicle or dirt bike use, or other threats.

(d) For species that are extirpated (or nearly so) in San Francisco, collect seed

from closest geographic populations in ecologically equivalent habitats, when

source populations are large enough not to be impaired by sampling of

approximately 50 to 100 seeds, or clones of approximately 50 individuals. 

Samples should be obtained randomly throughout the source population,

equalizing numbers of propagules (seeds, cuttings, etc.) from each plant

sampled. Plants should be propagated in native plant nurseries for

reintroduction or augmentation of San Francisco populations, following expert

scientific peer review of reintroduction/augmentation plans (including detailed,

specific taxonomic and ecological evaluation). Augmentation plans should

also include a provision for low-level dispersal (immigration) of seeds or

plants to habitats that have been fragmented by urban development, but still

contain small, isolated populations. 

(e) For target species that persist in San Francisco, apply management actions

to reduce the principal impediments to the persistence or re-expansion of

populations, such as invasive nonnative vegetation, adverse soil modifications

(e.g., irrigation, fertilization, leaf litter of nonnative vegetation), and excessive

trampling. Population augmentation by transplanting nursery-propagated stock

should be attempted only if effective habitat enhancement fails to promote

viable populations; planting should not be used as a primary method of

conserving remnant populations of target species.

(f) For rare target species with chronically or periodically small populations,

maintain either stored seed, or pedigreed clones in cultivation, as appropriate

for the life-history of the species.

(g) Conduct research on species-specific techniques for propagation,

transplanting and establishment techniques, microenvironmental requirements,

and species associations (competitor, mutualist, pollinator relationships).
3 Recovery action priority is defined at the beginning of the Implementation

Schedule on page 205.

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V. STEPDOWN NARRATIVE OF RECOVERY ACTIONS

A. San Francisco Lessingia

A 1. Protect, maintain, and enhance existing populations and habitat.

A 1.1. Protect and manage populations on the Presidio

A 1.1.1. Develop and implement general and specific management

plans. Work with the Golden Gate National Recreation Area and the

Presidio Trust to complete Recovery Actions A 1.1.1.1 through A

1.1.1.4 below.

A 1.1.1.1. Develop and implement site-specific implementation

plans under the Presidio Vegetation Management Plan to

prescribe maintenance and enhancement activities consistent

with this recovery plan for all lessingia sites of the Presidio. 

Site-specific actions under this action may require section 7

consultation on the proposed site-specific actions, including

management of the buffer areas around structures. (Priority 1)3

A 1.1.1.2. Delineate and permanently dedicate Golden Gate

National Recreation Area and Presidio Trust lands identified in

this recovery plan (including prospective restoration areas) to

management of dune vegetation supportive of San Francisco

lessingia. (Priority 1)

A 1.1.1.3. Evaluate existing site-specific management plans,

and develop and implement necessary amendments, including

monitoring, for the Lobos Dunes, Battery Caulfield, Public

Health Services Hospital, Rob Hill, Upper Baker Beach Dunes,
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Presidio Golf Course Site, and Wherry Dunes populations of

San Francisco lessingia.

A 1.1.1.3.1 In coordination with the National Park

Service, develop a management plan for the two portions

of the Special Management Zone designated in the

Presidio Vegetation Management Plan that overlap with

the Presidio Recovery Unit. Determine what changes to

the management of the historic forest area within these

blocks, if any, are needed to meet the need for

establishing a larger, continuous population of San

Francisco lessingia with connection between the Lobos

Dunes population and the contiguous areas to the north. 

These two Special Management Zone areas include all

or portions of the Lobos Dunes Site and the Upper Baker

Beach Dunes. (Priority 1)

A 1.1.1.3.2 Evaluate the West Washington Housing

Area to determine its suitability for addition to the

Presidio Recovery Unit. The Presidio Trust has

proposed that restoration of dune scrub habitat to enlarge

the Presidio Recovery Unit following the removal of

West Washington Housing over the next 30 years may

increase the likelihood of recovery for San Francisco

lessingia. This area is located adjacent to both the

Wherry Housing and Rob Hill Reserves. The suitability

of the site and potential to expand the currently proposed

reserves should be evaluated and incorporated into the

recovery plan, if appropriate. (Priority 2)

A 1.1.1.3.3 Develop and implement site-specific

implementation plan for the Building 1750 and Bowley

Street Interim Management Area. The National Park

Service and the Presidio Trust will work with us to

develop and implement a site-specific management plan. 

During and after implementation of restoration of the
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lands downwind from the Building 1750 and Bowley

Street Interim Management Area (see Figure 11),

ecosystem function will be evaluated. If ecosystem

function, including wind scouring, weed control,

moisture balance, and San Francisco lessingia

recruitment appears adequate to meet recovery

objectives, then we recommend continuing in perpetuity

the interim forest management practices (replacement

with lower stature trees, lower tree densities, higher tree

canopy floor, etc.) described in the Presidio Vegetation

Management Plan. If aspects of ecosystem function are

not adequate to meet recovery objectives, in

coordination with the Presidio Trust and Golden Gate

National Recreation Area, develop additional measures

to improve ecosystem function and reevaluate effects of

revised management. (Priority 1)

A 1.1.1.3.4 Evaluate existing site-specific management

plans, and develop and implement necessary

amendments for the rest of the sites, including

monitoring, for the Battery Caulfield, Public Health

Services Hospital, Rob Hill, and Wherry Dunes

populations of San Francisco lessingia. (Priority 1)

A 1.1.1.4. Commit timely and adequate resources to implement

vegetation management plans affecting San Francisco lessingia,

sufficient to achieve interim recovery criteria. Reduce invasive

nonnative vegetation to low and insignificant levels, or

eradicate them completely if possible, at all San Francisco

lessingia sites. (Priority 1)

The management plans should be developed collaboratively

among Golden Gate National Recreation Area, The Presidio

Trust, California Department of Fish and Game, and the U.S.

Fish and Wildlife Service, with public outreach, review,

comment, and lead agency response to comments. Management
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plans should comply with the recovery strategy outlined above,

and should apply to both Golden Gate National Recreation Area

and Presidio Trust lands that are proposed to support San

Francisco lessingia in the future. Plans should address

vegetation management techniques (e.g., herbicide use, manual

weed removal, managed or semi-natural disturbance regimes,

trampling intensity, public access and rotational trail closures,

and experimental modification of established policies). [Please

note that these site-specific vegetation management plans are

referred to by the National Park Service as restoration action

plans, and are intended to be sufficient to guide on-the-ground

work. The existing Presidio Vegetation Management Plan is a 

more strategic level planning document.]

A 1.1.2. Monitor all Presidio populations. Monitoring of San

Francisco lessingia populations should be integrated into management

plans (Recovery Action A 1.1.1). Monitoring should be conducted

indefinitely on an annual basis until recovery criteria are met for the

Presidio Recovery Unit. After the criteria are met, results will be

evaluated to determine the frequency and intensity of further

monitoring. Monitoring methods should include: periodic false-color

infrared vertical aerial photographs of each site; quantitative stratified

sampling of vegetation; annual census of mature plants; some

population sampling and analysis of reproductive output. Other

methods may be determined during development of management and

monitoring plans to be appropriate. These methods need to be

sufficient to estimate population numbers, define habitat structure, and

characterize vegetation cover and species composition necessary to

evaluate progress toward meeting the recovery criteria.. Use of these

other methods must be approved by the U.S. Fish and Wildlife Service

and peer reviewed. Monitoring should also focus on problem areas for

invasive nonnative plants. (Priority 2)

A 1.1.3. Establish permanent and seasonal staff dedicated to lessingia

vegetation management. Establish and fund a permanent full-time

vegetation technician to implement and oversee maintenance (and
185

restoration) at all Presidio populations. The position should be funded

at least until recovery criteria for the Presidio Recovery Unit are met. 

This technician should coordinate efforts with analogous dune

management activities at Fort Funston. Establish seasonal vegetation

technical positions in support of the full-time technician as needed.

(Priority 1)

A 1.2. Protect the Daly City population.

A 1.2.1. Protect the Daly City population under its existing ownership. 

Establish interim or permanent cooperative agreements, or conservation

easements, with the Daly City site’s private and municipal landowners

to protect the site against adverse land uses, and to enable

implementation of beneficial maintenance and enhancement activities

(principally control of invasive nonnative vegetation). (Priority 1)

A 1.2.2. Acquire Daly City parcels within proposed San Francisco

lessingia reserve. Acquire Daly City parcels supporting San Francisco

lessingia, associated species, as well as restorable habitat on the sand

slopes above Hillside Park and Bonnie Street, to ensure permanent

protection and ability to manage the site without conflicts of land use.

(Priority 1)

A 1.2.3. Establish a land manager and endowment fund for site

management. Select a permanent land manager and provide an

endowment to maintain and manage the site. Maintenance and

management obligations would include control, progressive removal

and reduction of nonnative vegetation, erosion control, regulation of

pedestrian access, public education and nature interpretation,

population monitoring of San Francisco lessingia, and our coordination

with neighbors. (Priority 2)

A 1.2.4. Prepare and implement a long-term management and

restoration plan. Prepare a site-specific management plan according to

the general management and (limited) restoration prescriptions of this

recovery plan. The content of the plan should correspond with that of
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the Presidio specific plans. (Recovery Actions A 1.1.1.3.1 – 1.1.1.3.4). 

Implement the plan in perpetuity. (Priority 2)

A 2. Restore habitat and reintroduce San Francisco lessingia at designated reserves.

A.2.1 Prepare plans for San Francisco lessingia reintroduction and genetics

management.

A 2.1.1 Prepare plans for habitat restoration and augmentation or

reintroduction of populations. Prepare comprehensive restoration plans

for San Francisco lessingia and reintroduction or augmentation plans at

the reserves specified below. Planning for Presidio sites that are

proposed for consolidation with the Lobos dunes reserve should be

prepared jointly, so these sites will form a unified, phased complex,

rather than independent units, each with its own plan. Plans should

implement general conceptual designs. For example, plans should

specify tasks for seed bank assessments of weeds and native species;

native plant/seed bank recruitment salvage where appropriate;

nonnative tree and iceplant (Carpobrotus sp.) removal; duff and debris

removal (including removal of organically enriched surface soil layers);

weed seed bank exhaustion; topographic recontouring; temporary postgrading dune stabilization; composition, pattern and density of local

native species transplanting; pattern and density of San Francisco

lessingia seeding; and contingency measures (including options for

delay of reintroduction) in case of drought or catastrophic failures.

(Priority 1)

A 2.1.2 Prepare a genetics management plan for San Francisco

lessingia. A genetics management plan should be prepared to

determine whether populations of San Francisco lessingia at Sunset

Heights in the Satellite Recovery Unit should be re-established using

Presidio or Daly City stock, or some mixture of the two resulting from

crossing off-site or by planting both types in the Recovery Units to

allow selection on site. The distinct genetic types of San Francisco

lessingia at the Presidio and Daly City populations require that we

analyze the results we could expect from crossing to determine the best
187

choice for re-establishment of populations at Sunset Heights. Our

policy requires the preparation of a genetics management plan that

incorporates the elements of the guidelines for propagation of the

Center for Plant Conservation (U.S. Fish and Wildlife Service 2000). 

This should happen prior to any planting in the Recovery Units. 

(Priority 1)

A 2.2. Expand populations and restore habitat at designated San Francisco

lessingia reserves.

A 2.2.1. Expand population and restore habitat above Lobos dunes site,

north and east of Lobos Creek to Battery Caulfield Road. Remove all

Monterey cypress (Cupressus macrocarpa) and iceplant on dune slopes

above (north and east) of Lobos Creek, to Battery Caulfield Road

(linking the San Francisco lessingia relict population there), prepare

and recontour slopes of dune sand surface, exhaust invasive weed

seed/bulb banks, and reintroduce native dune scrub and grassland

vegetation with San Francisco lessingia, in patchy, low density initial

populations. Establish a consolidated, expanded Lobos Dune

population of San Francisco lessingia, according to the site- specific

implementation plan for this reserve (Recovery Action A 2.1.1). 

Portions of this area are within the Special Management Zone defined

in the Presidio Vegetation Management Plan and will be subject to

additional site-specific implementation planning efforts by the

National Park Service and U.S. Fish and Wildlife Service under action

A.1.1.1.3.1. (Priority 1)

A 2.2.2. Expand populations and restore habitat from Lobos Dune site

to Baker Beach dunes. Incrementally remove the Monterey pine (Pinus

radiata) and Monterey cypress grove from the north end of Baker

Beach, including removal of duff/leaf litter removal (but no

recontouring), following the restoration plan for this site (treated as part

of the Wherry site). These actions will open wind fetch for the Wherry

Dunes and open habitat for dune scrub supporting San Francisco

lessingia. Establish a roadside buffer zone of transplanted native

woody dune scrub vegetation, but allow natural recolonization by
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native dune scrub, along with active suppression of invasion by

nonnative species there; or establish only patchy distribution of native

dune scrub vegetation by limited transplanting of locally propagated

populations if natural recruitment is inadequate. Establish multiple

founder colonies of San Francisco lessingia at the same time as pioneer

species are being extirpated. Remove nonnative conifers from upper

dune slopes near Lincoln Boulevard at the south end of Baker Beach to

re-open wind fetch for expanded Lobos Dune habitat. Establish a

consolidated, expanded Lobos dune population of San Francisco

lessingia with higher long-term viability, according to the plan for this

site (Recovery Action A 2.1). Portions of this area are within the

Special Management Zone defined in the Presidio Vegetation

Management Plan and will be subject to additional planning efforts by

the National Park Service and U.S. Fish and Wildlife Service under

action A 1.1.1.3.1. (Priority 1)

A 2.2.3. Expand populations and restore habitat from Lobos Dunes

Reserve through Wherry Housing area (Lobos/Wherry/Baker Reserve). 

Begin the previously-planned phased demolition of Wherry Housing,

and implement the restoration/ reintroduction plan for this portion of

the Presidio Recovery Unit. Establish a consolidated, expanded Lobos

dune population of San Francisco lessingia with higher long-term

viability, according to the plan for this complex (Recovery Action A

2.1). Portions of this area are within the Special Management Zone

defined in the Presidio Vegetation Management Plan and will be

subject to additional planning efforts by the National Park Service and

U.S. Fish and Wildlife Service under action A 1.1.1.3.1. (Priority 2)

A 2.2.4. Expand populations and restore habitat at the Public Health

Services Hospital lessingia Reserve. Remove the Monterey pine and

iceplant understory north of the Public Health Services Hospital, and

remove duff/leaf litter layer, following the restoration plan for this site. 

Reintroduce sparse dune scrub and grassland vegetation with San

Francisco lessingia according to the plan for this reserve. (Priority 2).
189

A 2.2.5. Expand populations and locally restore/enhance habitat within

the Daly City lessingia Reserve for San Francisco lessingia. 

Incrementally eradicate iceplant and annual nonnative grasses on the

Hillside Park sand slopes, following the restoration plan for this

reserve. To prevent excessive slope destabilization, work upslope in

increments, and facilitate natural recolonization by dune scrub

vegetation, or transplant locally propagated natives. Establish a mosaic

of bare patches in slopes dominated by nonnative vegetation (such as

horseweed [Conyza canadensis], annual grasses), and suppress

nonnative vegetation while establishing a new matrix of dune scrub

vegetation around bare patches. Sow a mix of native annual seed

including local San Francisco lessingia in bare patches. (Priority 2)

A 2.2.6. Restore habitat and reintroduce San Francisco lessingia to

dunes at Fort Funston. Incrementally eradicate iceplant and nonnative

conifers and scrub at Fort Funston (Golden Gate National Recreation

Area) and reintroduce San Francisco lessingia from Daly City

according to the plan for this site (Recovery Action A 2.1). 

Development of a restoration and reintroduction plan for Fort Funston

should incorporate results of site-specific studies to determine

recreation use management consistent with maintaining a dune scrub

community and meeting the recovery criteria for San Francisco

lessingia (see Recovery Action A 4.5). Plans must also define buffers

and buffer management around historic structures. (Priority 2)

A 2.3. Establish beach layia (Layia carnosa) to the Fort Funston lessingia

Reserve for San Francisco lessingia. Integrate establishment of beach layia

(federally listed as endangered) with lessingia reserve at Fort Funston,

consistent with its recovery plan (U.S. Fish and Wildlife Service 1998b).

Beach layia seed source should be the core population from Point Reyes.

(Priority 2)

A 2.4. Establish new San Francisco lessingia population(s) at restored satellite

reserve dune remnants.
190

A 2.4.1. Secure the Hawk Hill site dune remnant in Sunset Heights. 

Ensure the Hawk Hill site, recently acquired by San Francisco City

Parks, is managed as a reserve for rare dune species. Establish an

endowment fund that will generate enough interest to support long-term

vegetation management to prevent reinvasion of dominant nonnative

vegetation. (Priority 2)

A 2.4.2. Secure privately owned portions of sand slopes and dunes

near Sutro Heights that are needed for this satellite reserve. Acquire

from willing sellers or establish conservation easements on the

privately owned portions of the sand slopes near Balboa Street and 48th

Avenue or other sites where opportunities for recovery may be found. 

Conservation easements must enable management of the sites as San

Francisco lessingia reserves. If acquired by fee-title, transfer title to the

City of San Francisco or other conservation organization, with deed

restrictions ensuring its conservation as a reserve for rare dune species. 

Establish an endowment fund that will generate enough interest to

support long-term vegetation management to prevent reinvasion of

dominant nonnative vegetation. (Priority 2)

A 2.4.3. Control or eradicate invasive nonnative vegetation within

satellite reserves. Incrementally eradicate iceplant, jubata grass

(Cortaderia jubata), ripgut brome (Bromus diandrus), and other

invasive nonnative plants at Hawk Hill and Sutro Heights reserves. 

Remove Monterey cypress trees and veldtgrass (Ehrharta erecta)

understory at Grandview Park. Expand margins of Sunset Heights Park

dune remnant by cutting back nonnative shrubs and trees along its

periphery and removing invasive species. Remove acacias, plume

acacia (Albizia lophanta), cypress, and iceplant from suitable sites in

the Sutro Heights area including the Cliff House sand slopes, and install

appropriate native buffer scrub vegetation around residential areas

there. (Priority 2)

A 2.4.4. Establish San Francisco lessingia in satellite reserves. In

areas where nonnative invasive vegetation has been cleared, establish

founder populations of San Francisco lessingia at Hawk Hill and
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suitable sites in the Sutro Heights area including the Cliff House sand

slopes, and at least one of the two other remnant dune habitat sites. 

Manage vegetation to a condition in which San Francisco lessingia

spontaneously regenerates and colonizes. (Priority 2)

A 3. Conduct public outreach, education, and coordination.

A 3.1. Prepare information presentations for public outreach. Prepare

scientifically accurate graphic, written, and oral accounts of the needs for

species recovery, the needs for habitat restoration, and the time and spatial

scales involved. Use endorsements (written statements of official support) of

restoration/reintroduction plans by government natural resource agencies and

nonprofit conservation groups for dissemination and information packages to

media, neighbors of reserves, and other interested public citizens. Present

information through public meetings, news media, and mailings. (Priority 1)

A 3.2. Conduct public outreach meetings prior to implementation of major

recovery actions. In advance of any tree removal, large-scale vegetation

management, or reintroduction actions, engage in substantial promotional

public outreach programs using local media, formal public notice, and public

meetings. Use media materials from Recovery Action A 3.1. (Priority 1)

A 3.3. Publicize and promote restoration sites prior to conspicuous site

modifications. Well in advance of site restoration actions, place informational

signs (or brochure dispensers) around restoration sites explaining prospective

actions and results. Include visual displays (photographic, diagrammatic) in

signs. (Priority 1)

A 3.4. Publicize project effects on scenic views and recreational use of the

Golden Gate and Presidio. Provide specific written, visual, and media

accounts of restoration projects’ compatibility (or incompatibility) with

recreational uses of these areas where restoration/reintroduction plans are

perceived to threaten established or locally popular recreational or esthetic land

uses (e.g., public access, dog walking at Fort Funston; forested viewsheds of

northern Presidio). Provide interpretive visual displays of scenic values of

viewsheds of Golden Gate, comparing views under forested and restored dune
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(tree removal) conditions. Provide explanatory accounts of temporary

exclosures during transplanting/establishment phases of restoration, and extent

of long-term public access. (Priority 2)

A 3.5. Actively support public volunteer participation. Promote and provide

Federal support (staff, funds) for public volunteer participation in monitoring

and site stewardship (vegetation management) activities, in coordination with

local nonprofit conservation organizations, particularly ones with volunteer site

stewardship programs in progress. (Priority 1)

A 3.6 Coordinate all outreach developed in actions 3.1 through 3.5 with

partners. Coordinate and make implementation decisions with partners to

determine appropriate implementation strategies, such as methods, phasing,

and prioritization. (See action A 6. for Participation Plan development.) After

implementation strategies are developed, work with partners to prepare and

carry out an outreach plan to ensure stakeholder input is heard and addressed. 

Partners should assess the appropriate types of outreach to inform and involve

the public and stakeholders most effectively. Outreach should inform the

public of the importance of endangered species recovery and the missions of

the National Park Service, U.S. Fish and Wildlife Service, and Presidio Trust,

to preserve the nation’s trust resources for present and future generations. 

Public information and outreach efforts can include, but not be limited to,

brochures, flyers, on-site signage, website updates, notices to mailing list of

stakeholders, media releases and interviews, and public meetings, public

walks, and volunteer participation. Public information will be disseminated

during, but not limited to, the following stages of restoration: prior to

implementation of major recovery actions, such as large-scale tree removal

(anything more than removing a handful of dead trees), on-site restoration

actions, or potential site closures. (Priority 1)

A 4. Conduct research in support of San Francisco lessingia recovery. San Francisco

lessingia habitat restoration, including both deliberate and accidental reintroduction

actions in the Presidio, began before the species was listed. Before the results were

available from scientific studies of the species’ life-history and competitive

interactions, restoration efforts were based on professional judgement of habitat needs

and observation of dispersal. These efforts were nonetheless extremely successful, at
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least in the short-term. Additional research may improve the long-term prospects for

success, but such research is not a prerequisite for initiating additional restoration and

reintroduction actions in the Presidio and Southern Recovery Units, nor is research

needed for initial dune scrub restoration in the Satellite Recovery Unit. These

conclusions are based on the practical, empirical results of the pilot projects at Lobos

Dunes, and the success of the unplanned introduction at Wherry Housing. It may be

necessary to study crosses between Presidio and Daly City types. The need is to be

determined by the genetics management plan specified in Recovery Action A.2.1.1. 

Additional research should be aimed at refining restoration and reintroduction

techniques, controlling nonnatives, establishment of suitable dune topography and

mobility, and critical aspects of the species’ ecology and taxonomy.

A 4.1. Study the usefulness of crosses between Presidio and Daly City genetic

types if this is determined necessary by the genetics management plan. 

(Priority 1)

A 4.2. Study variation in ecological and genetic characteristics of San

Francisco lessingia. Study patterns of population variation in significant

ecological traits among and within relict populations, and compare with

reintroduced populations, including both field and common-garden

approaches. Determine whether significant genetic differentiation occurs

within or between San Francisco lessingia populations or colonies, and assess

patterns of genetic diversity. Study the long term viability of seed stored under

ideal (refrigerated) conditions. Use the results to guide decisions about seed

sources for satellite reserves, controlled mixing of artificially isolated

populations in the Presidio, and the need to continue collection of 500 seeds

per year under Recovery Action A 5.2. (Priority 3)

A 4.3. Study rates and modes of recolonization by invasive nonnative plants,

and control methods. Study rates and modes of recolonization by invasive

nonnative plants (Appendix III) and methods of control, including seasonal

variation in all applied techniques. Techniques to be studied should include

manual removal, applications of synthetic herbicides appropriate for use in

parks (including glyphosate and grass-specific herbicides), natural foliar

desiccants (e.g., vinegar), and in some locations, earthmoving equipment

(scraping, burial). The study plans developed for this action should specify the
194

range of techniques tested and should include evaluation of any additional

techniques determined to be appropriate for park use and for the particular

species and habitats present on the study sites. (Priority 2)

A 4.4. Study the pollination ecology of San Francisco lessingia. Determine

the mode and vectors of pollination that contribute most to seed production.

(Priority 3)

A 4.5. Experimentally investigate feasibility of rotational trail closure /

temporary pedestrian exclosure as vegetation management technique. Study

rotational closure and opening of pedestrian trails and exclosures to determine

the potential efficacy of intermittent (among years) seasonally timed intensive

trampling as a method of maintaining disturbed semi-open dune vegetation

favorable to native annual dune plant species, including San Francisco

lessingia, at locations with high demand for recreational access (e.g., Fort

Funston).

After re-establishment of San Francisco lessingia and other federally listed

species and plant species of concern that occur in dune scrub habitats,

especially beach layia, begin study for the Fort Funston habitat. Include study

areas to observe the effects of no management, closure, and rotational use. 

Please see the actions for re-establishment of sensitive plant species. Related

actions include: A 1.1.1.4, A 1.1.3, A 2.2.6, A 2.3, A 4., A 4.3, A 4.5, A 4.6, A

4.7, and A 4.8. (Priority 1)

A 4.6. Conduct field experiments on restoring and anthropomorphically

managing dune topography. Conduct field experiments on managing and

restoring dune topography and mobility within the land management

constraints of the Presidio and Fort Funston, aimed at establishing undulating

to steep dune topography and mosaics of persistent, limited blowout activity

that are suitable for San Francisco lessingia regeneration and colonization. 

Determine rates of erosion and deposition at spatial scales favorable to the

species under field conditions. (Priority 2)
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A 4.7. Study seed dispersal and seed bank ecology of San Francisco lessingia. 

Study recruitment of different age-classes of seeds, longevity of dormant seeds,

and seed dispersal. (Priority 3)

A 4.8. Study effects of substrate conditions on growth and reproduction of San

Francisco lessingia. Compare growth and reproduction on natural variation in

local substrates (e.g., young dune, old dune, Colma Formation sand). Study

effects of sand mineral nutrition, mycorrhizae, slope aspect, microclimate),

comparing differences between Presidio and Daly City populations. (Priority 2)

A 4.9. Clarify taxonomic relationships between San Francisco lessingia and

allied species. Assess the biosystematic relationship between San Francisco

lessingia and similar species (e.g., Lessingia glandulifera varieties).

(Priority 3)

A 5. Offsite conservation measures.

A 5.1. Establish cultivated populations. Establish cultivated populations of

San Francisco lessingia at regional botanical gardens for study (providing

sources of seed and herbarium specimens for research, alleviating some need

for harvest of wild populations) and public education. (Priority 2)

A 5.2. Maintain stored seed bank to ensure survival in case of population

failure in reserves. Collect seed from the Daly City and Presidio population

groups annually for deposit in seed banks certified by the Center for Plant

Conservation, as well as for distribution to local conservators approved by us

(e.g., Golden Gate National Recreation Area, Strybing Arboretum). Collect

about 25 capitula (seedheads, each with about 20 seeds each) from source

populations in each of the Presidio and Southern Recovery Units to get 500

seeds from each Recovery Unit. Periodically test lots of old seed for

germinability. Determine whether 500 per year per group are needed to

maintain 1,000 viable seed. After 5 years, adjust annual collection rate as

indicated by seed survival statistics. (Priority 3)

A 6. Develop a participation plan with all partners and stakeholders that outlines a

strategy to implement recovery actions. Our 1994 Interagency Cooperative Policy on
196

Recovery Plan Participation and Implementation Under the Endangered Species Act

(U.S. Fish and Wildlife Service and National Oceanic and Atmospheric

Administration 1994a) provides for a participation plan process, which involves all

appropriate agencies and affected interests in a mutually-developed strategy to

implement recovery actions. A participation plan for implementing recovery actions

for San Francisco lessingia that includes all partners should be developed. In addition

to outlining a strategy to implement recovery actions, the participation plan should

include strategies for evaluation of progress and needs for plan revision. Participation

plans may also achieve the policy’s goal of providing for timely recovery of species

while minimizing social and economic impacts. (Priority 1)

B. Raven’s Manzanita

B 1. Protect, maintain, and enhance the relict natural clone of Raven’s manzanita at

the World War II Memorial, Presidio.

B 1.1. Control nonnative vegetation. Suppress nonnative invasive plants in

serpentine grassland and scrub around the natural clone of Raven’s manzanita.

Aim at local eradication of nonnative vegetation, but achieve at least

suppression to insignificant levels. Aim at keeping invasion front at least 30

meters (100 feet) away from Raven’s manzanita. Prohibit use of any

herbicides that affect broadleaf plants within 30 meters (100 feet) of the

manzanita to avoid any possible contact with herbicide drift. (Priority 1)

B 1.2. Monitor growth, reproduction, and clone size of Raven’s manzanita

annually. Monitor flower production, timing, insect visitors to flowers, seed

set, and seed dispersal. Monitor growth at areas of contact with above ground

parts of competing shrubs. (Priority 2)

B 1.3. Manage native vegetation. Based on monitoring of growth of Raven’s

manzanita, prune or remove competing native vegetation if it causes significant

depression of growth of portions of the original clone or daughter clones.

(Priority 1)

B 2. Protect, maintain, enhance, and increase transplanted Raven’s manzanita

populations in the Presidio.
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B 2.1. Monitor growth and clone size of established transplants annually. The

size of plants, rate of growth, and proportions of live and necrotic patches

(blight-induced dieback) should be monitored at least once annually, and

preferably at the beginning, peak, and end of each growing season. Production

of flowers and fruit should be monitored annually. Fruit should be sampled

annually to estimate production of viable seed. (Priority 2)

B 2.2. Control nonnative vegetation. Suppress nonnative invasive plants, and

native shrubs, where necessary (see Recovery Action B 1.1), at sites of

previous transplants along Lincoln Boulevard and Inspiration Point. Prohibit

use of herbicides that affect broadleaf species within 30 meters (100 feet) of

these sites to prevent damage by herbicide drift or accidental application.

(Priority 1)

B 2.3. Survey and select additional serpentine outcrop sites for further

transplantation of clones within the Presidio. Selected sites should include

serpentine bedrock outcrops in relatively stable positions in the Presidio bluffs,

and at least one additional location away from the bluffs. Criteria for selection

include sparse vegetation, a lack of strong local invasion pressure by nonnative

herbaceous vegetation, and absence of federally listed species or species of

concern. Preference for reintroduction sites should be given to serpentine

outcrops that lack significant native serpentine plant populations. (Priority 2)

B 2.4. Prepare sites as necessary for transplantation of propagated clones in

the Presidio. To prepare sites, remove any nonnative trees within peak seed

dispersal distance (all trees upwind to a distance of at least 90 meters [300

feet]) to avoid recolonization, shading, and fog drip effects. Remove all

invasive nonnative shrubs and herbaceous species (particularly myoporum

[Myoporum laetum], iceplant, plume acacia, acacia [Acacia sp.], and jubata

grass) within at least 300 meters (1,000 feet) upwind of the manzanita site; for

jubata grass, a species with small plumed seeds that are highly capable of longdistance dispersal, all plants upwind should be eradicated). For sites restored

under former tree cover, reintroduce native associated serpentine-affinity

species from local sources. (Priority 2)
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B 2.5. Transplant replicated clones at selected prepared sites. Transplant

Presidio clones at selected reintroduction sites. Plant in groups to allow for

mortality, and to increase attractiveness of the population to pollinators. If

possible, include new clones propagated from seed from the original plant, or

from controlled self-pollinated Presidio clones. Provide frequent monitoring

(weekly) and intensive post-transplant care (including supplemental watering,

mycorrhizal inoculation, and low-level application of nutrients, especially

organic nitrogen) for one growing season, or two seasons if drought conditions

prevail. At least 3 additional populations remote from the parent colony

should be established, consisting of at least 10 replicate clones that survive and

grow vigorously at least 5 years after transplanting. (Priority 2)

B 2.6. Monitor transplanted replicated clones. After establishment (beginning

third growing season), monitor growth and survivorship of new transplanted

clones at least four times annually. Shift to minimum annual monitoring after

the fifth growing season. Monitor associated vegetation. Provide adaptive

levels of vegetation management in response to nonnative species invasion. If

sites fail due to interference from excessive growth of native serpentine

grassland or scrub species, abandon the colony and replicate two additional

reintroduction sites elsewhere in the Presidio (following Recovery Actions

B.2.1 to B 2.6). (Priority 2)

B 2.7. Identify and protect known and potential serpentine outcrop sites. 

Survey and permanently protect all serpentine outcrops and near-surface

occurrences of serpentine in the Presidio against development to reserve them

for future restoration and reintroduction. All serpentine sites should be

managed by Golden Gate National Recreation Area natural resources staff, and

should eventually be transferred to Golden Gate National Recreation Area.

(Priority 2)

B 2.8. Reintroduce associated serpentine plant species. Incorporate

reintroduction of other native serpentine grassland species in decline (or

extirpated) in San Francisco. Include plans to reintroduce other endangered

species, consistent with an approved recovery plan (U.S. Fish and Wildlife

Service 1998a). Select stock from local San Francisco sources or closest

serpentine habitats. (Priority 3)
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B 3. Restore habitat and establish new populations in San Francisco outside the

Presidio.

B 3.1. Evaluate and select candidate reserve sites for reintroduction around

San Francisco. Priority should be given to bedrock outcrops within the Fort

Point-Hunter’s Point serpentine belt (Figure 3). The next highest priority

should be greenstone (particularly basalts) outcrops. Lowest preference should

be given to other Franciscan rocks lacking mafic or ultramafic chemistry, but

some should be included in the final array of selected sites. Preference should

also be given to sites that include areas with species-poor native plant

communities owing to past disturbance, to avoid impacts to relict native

vegetation. Selected sites collectively should exhibit variation in slope, aspect,

fog sheltering, and wind exposure. A total of at least five sites should be

selected. (Priority 2)

B 3.2. Secure selected reserve sites. Acquire sites through fee title purchase

with conservation deed restrictions, or secure with conservation easements, or

establish cooperative agreements for reintroduction sites that are municipally

owned. Agreements and easements must cover the full scope of actions

prescribed in restoration and recovery plans (Recovery Action B 3.3).

(Priority 2)

B 3.3. Prepare restoration and reintroduction plans. Prepare restoration and

reintroduction plans for all sites (conceptual general plan) and for specific sites

(implementation plans), with emphasis on removal and suppression of

regeneration of nonnative vegetation. At least three interior San Francisco

sites should include plans for mixed transplanted populations of both San

Franciscan manzanitas. At least 30 replicate transplants of Raven’s manzanita

should be assigned each site to allow for mortality. Plans should also include

provisions for whatever removal of nonnative trees and shrubs is needed,

eradication of invasive nonnative vegetation (and maintenance), protection of

native vegetation, post-transplant care and maintenance, and reintroduction of

associated native species of native grassland and coastal scrub communities,

chosen to fit local substrate conditions, and derived from appropriate source

populations. (Priority 2)
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B 3.4. Propagate stock for founder populations. Propagate Raven’s manzanita

and Franciscan manzanita stock for transplanting, using clones of seedpropagated, genetically controlled plants. Priorities for genetic management of

stock should be based on recommendations of a scientific peer review panel

(see Recovery Action B 5.7, below). Part of the propagation process is to

inoculate stock with mycorrhizae (fungi that associate with roots) derived from

the remnant Presidio population. (Priority 2)

B 3.5. Prepare reserve sites for reintroduction. Prepare sites for reintroduction

according to restoration and reintroduction plans by removing and suppressing

regeneration of nonnative vegetation, but only after full completion of public

outreach actions. (Priority 2)

B 3.6. Transplant and establish propagated founder plants. Transplant and

perform post-transplant maintenance and intensive monitoring using the same

methods as for transplanted Presidio populations (Recovery Actions B 2.5 to B

2.6). (Priority 2)

B 3.7. Monitor transplants. Monitor transplants and subsequent generations as

for transplanted Presidio populations (Recovery Actions B 1.2, B 2.6). If sites

fail due to interference from excessive growth of native coastal scrub or

grassland species, abandon the colony and replicate two additional

reintroduction sites elsewhere on site or at new sites. (Priority 3)

B 4. Public outreach, education, and coordination.

Public outreach and education are particularly important for Raven’s manzanita

recovery because recovery depends on habitat restoration and reintroduction in urban

reserves. Habitat restoration in San Francisco may involve removal of ornamental

nonnative species. Additionally, reintroduction of endangered species may cause

neighbors to have concerns about land use restrictions. Community support, including

support of local governments, will be essential to implementation of recovery actions.

B 4.1. Promote native manzanita recovery broadly through information and

outreach efforts. Promote public outreach efforts about native endemic

manzanitas and their historic habitats to develop support for experimental
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restoration and reintroduction projects, as well as conservation of the remnant

Presidio population. We should collaborate with local natural history

museums, nonprofit conservation organizations, Golden Gate National

Recreation Area, Presidio Trust, local schools (including colleges and

universities), and local botanical gardens to conduct informational programs

(speakers, slide talks, field trips). (Priority 2)

B 4.2. Promote native manzanita recovery at the local community level. Well

in advance of restoration activities – especially before removing any trees or

conducting large-scale vegetation management or reintroduction actions,

conduct promotional public outreach to nearby neighborhoods through

neighborhood associations, park associations, and local schools. It also is

important to conduct broader public outreach programs using local media,

formal public notice, and public meetings. Such outreach should be done in

coordination with our office. Prepare scientifically accurate, written and oral

accounts and graphic displays explaining the needs for species recovery, the

needs for habitat restoration, and the time and spatial scales involved. Compile

endorsements (written statements of official support) from government natural

resource agencies and nonprofit conservation groups regarding

restoration/reintroduction plans. Develop local support by using endorsements

in information packages for the media, neighbors of prospective reserves, and

other interested public citizens. Present accounts through public meetings,

news media, and mailings. (Priority 2)

B 4.3. Promote native manzanita recovery through stewardship. Promote and

provide our support (staff, funds) to public volunteer participation in

monitoring and site stewardship (vegetation management) activities, in

coordination with local nonprofit, volunteer-based conservation organizations,

particularly ones with established volunteer site stewardship programs.

(Priority 3) 

B 4.4. Establish public demonstration gardens displaying cultivated Raven’s

manzanita clones. At or near potential sites of reintroduction, maintain

cultivated clones of Raven’s manzanita and selected associated species in

neighborhood public demonstration gardens, in cooperation with local

nonprofit urban horticulture and conservation organizations, in advance of
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restoration and reintroduction actions. Establish informational and stewardship

programs for demonstration gardens to develop local community support for

restoration and reintroduction actions. 

(Priority 3)

B 5. Conduct research and development actions supportive of recovery.

B 5.1. Search for “Raven’s-like” Tamalpais manzanitas. Perform field

surveys for individuals of Tamalpais manzanita (Arctostaphylos montana)

from Marin County that are similar to Raven’s manzanita, or individuals with

at least multiple key characters (e.g., foliar, floral traits) overlapping with

Raven’s manzanita. Mark, map, collect, and propagate such specimens. 

Utilize propagated clones in studies of taxonomic relationships, or, if needed,

for introgressive breeding. (Priority 2)

B 5.2. Investigate seed production and propagation of Raven’s manzanita. 

Study seed production of artificially controlled self-pollinated Raven’s

manzanita. Determine whether viable seed can be produced through selfpollination. Compare with seed production of self-pollinated and outcrossed

manzanitas (Arctostaphylos montana, A. franciscana, local A. uva-ursi [San

Bruno, Point Reyes populations], and Monterey A. hookeri subspecies). 

Cultivate, label, and propagate all progeny, retaining pedigrees of each. 

Develop more reliable, rapid, and efficient techniques of clonal propagation of

seed-propagated plants. (Priority 2)

B 5.3. Investigate taxonomic status and relationships of Raven’s manzanita. 

Study in greater depth the systematic (including genetic) relationships between

Raven’s manzanita and other similar manzanita taxa of the central California

coast region (e.g., Arctostaphylos franciscana, A. montana, A. pungens, A.

hookeri, and A. uva-ursi, or other allied taxa). Molecular genetic analysis of a

wider range of taxa in the genus, morphometric comparisons, studies of

population variability, and synthesis of artificial hybrids should be addressed in

research. (Priority 2)

B 5.4. Investigate substrate and plant interactions with Raven’s manzanita. 

Study physiological ecology of substrate conditions (particularly variable
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mafic and ultramafic soil conditions) on growth, competition, and reproduction

of endemic San Francisco manzanita subspecies and dominant associated

native and nonnative species. Study potential effects of serpentine substrate on

biotic interactions such as herbivory, pathogen resistance, and mycorrhizal

function. (Priority 2)

B 5.5. Investigate reproductive ecology of Raven’s manzanita. Study

reproductive ecology of the remnant Presidio population (pollination, seed set,

seed dispersal, and seed germination and establishment), and perform

controlled experiments on germination and seedling growth responses of

cultivated seed to relevant environmental and biotic factors that can be

managed (e.g., fire, temperature, microbes, gut passage, etc.). (Priority 2)

B 5.6. Investigate practical specific nonnative vegetation control techniques. 

Study, develop, and refine effective and practical techniques of controlling or

eradicating invasive nonnative plants on steep, rocky slopes in local urban

settings, and on serpentine. (Priority 1)

B 5.7. Assemble a scientific review panel to evaluate and recommend a

genetic management program for Raven’s manzanita. An expert scientific peer

review panel should evaluate the appropriateness and need for a breeding

program for Raven’s manzanita. We, the Golden Gate National Recreation

Area (National Park Service), and the scientific panel should collaboratively

develop a genetic management plan. The plan should describe a breeding

program and recommendations on the use of newly bred Raven’s manzanita

plants in reintroduction. The panel should include experts in manzanita

biology, plant conservation genetics, plant breeding, and plant propagation.

(Priority 2) 

B 6. Undertake offsite conservation measures.

B 6.1. Collect and store seed of Raven’s manzanita. If self-pollinated Raven’s

manzanita seed are produced in the Presidio, approximately one third should be

collected for long-term storage, and be deposited at a botanical garden certified

by the Center for Plant Conservation. Additional seed should be stored by the 

Golden Gate National Recreation Area and local botanical gardens approved
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by us. One third of harvested seed should also be collected for testing for

hybridity, and propagation in support of experimental reintroduction if seed are

not hybrid (see Recovery Action B 3.2). (Priority 3)

B 6.2. Cultivate more Raven’s manzanita in botanical gardens. Raven’s

manzanita and Franciscan manzanita should be maintained in propagation and

in plantings at local botanical gardens approved by us. Informational signs

explaining Raven’s manzanita’s endangered status, rarity and endemism,

taxonomic status, and history of decline, should accompany plantings. 

Cultivated specimens rather than outplanted or wild populations should be used

to supply herbarium and research material. (Priority 3)

B 6.3. Promote and authorize appropriate informational and outreach use of

Raven’s manzanita. Cultivation of Raven’s manzanita for informational

purposes in San Francisco outside of botanical gardens should be encouraged

where it would be useful in: (1) increasing public support and appreciation of

this species; (2) encouraging volunteer participation in the stewardship of

reintroduction sites; and (3) reassuring neighbors of potential reintroduction

sites that the presence of this endangered plant species would not interfere with

reasonable enjoyment of public lands where it is reintroduced. Local sponsors

of demonstration plantings must comply any requirements for approvals by

landowners. Qualified local native plant propagators should be permitted by us

to propagate Raven’s manzanita from Federal land at the Presidio, using stock

from daughter clones, not the single original plant. It may be appropriate for

such propagation permits to include stipulations about the disposition of

propagated material. Demonstration plantings should be coordinated with

local nonprofit conservation and horticultural organizations. (Priority 3)
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VI. IMPLEMENTATION SCHEDULE

The implementation schedule that follows outlines actions and estimated costs for this

recovery plan. It is a guide for meeting the objectives discussed in Chapter IV of this

recovery plan. This schedule describes and prioritizes recovery actions, provides an

estimated time table for performance of recovery actions, indicates the responsible

agencies, and estimates costs of performing recovery actions. These actions when

accomplished should further the recovery and conservation of the covered species.

Key to Acronyms used in the Implementation Schedule

Definition of recovery action priorities:

Priority 1 – An action that must be taken to prevent extinction or prevent the

species from declining irreversibly in the foreseeable future.

Priority 2 – An action that must be taken to prevent a significant decline in

species population or habitat quality, or some other significant

negative impact short of extinction.

Priority 3 – All other actions necessary to meet recovery or conservation

objectives.

Definition of recovery action durations:

Continual – An action that will be implemented on a routine basis once

begun.

Ongoing – An action that is currently being implemented and will continue

until action is no longer necessary.

Unknown – Either action duration or associated costs are not known at this

time.
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Abbreviations:

TBD To be determined

Responsible parties:

CDFG California Department of Fish and Game

DC City of Daly City

GGNRA Golden Gate National Recreation Area, National Park Service

OWN Private landowners or parties

PC Private consultant (contractor)

PNP Private nonprofit organizations (local conservation, horticultural

organizations)

PT Presidio Trust

SF City of San Francisco Department of Parks and Recreation

USFWS U.S. Fish and Wildlife Service

UNIV undetermined university research laboratories 

* Primary responsible partner: a partner likely to take the lead on,

or have an especially large role in, implementing the recovery

action. 
Implementation Schedule for Northern San Francisco Peninsula Coastal Plants Recovery P

Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

1 A 1.1.1.1 Develop site-specific implementation

plans under the Presidio VegetationManagement Plan to prescribemaintenance and enhancementactivities consistent with this recoveryplan for San Francisco lessingia sitesat the Presidio. Evaluate and revisemanagement plans as needed.

 2 years *GGNRA

USFWS

PT, CDFG

3 1 2 0 

1 A 1.1.1.2 Delineate and permanently dedicate

Golden Gate National Recreation Areaand Presidio Trust lands for recoveryof San Francisco lessingia.

 3 years *GGNRA

USFWS

*PT, CDFG

3 2 0.5 0.5 

1 A 1.1.1.3.1 In coordination with the National Park

Service, develop a management planfor the two portions of the SpecialManagement Zone designated in thePresidio Vegetation Management Planthat overlap with the San Franciscolessingia Presidio Recovery Unit.

2 years *GGNRA, PT,

*USFWS

2.4 1.2 1.2 0 

1 A 1.1.1.3.3 Develop and implement site specific

implementation plan for the Building1750 and Bowley Street InterimManagement Area

30 *GGNRA, *PT,

*USFWS

7.2 1.4 0.2 0.2 

1 A 1.1.1.3.4 Evaluate other existing site-specific

management and monitoring plans forPresidio populations of San Franciscolessingia; amend as needed.

 ongoing *GGNRA

USFWS

*PT, CDFG

19.8 2 2.4 0.55 

207
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

1 A 1.1.1.4 Commit resources to implement

vegetation management andrestoration action plans affecting SanFrancisco lessingia at the Presidio.

ongoing *GGNRA

USFWS

*PT, CDFG

TBD – – – 

1 A 1.1.3 Establish permanent and seasonal staff

dedicated to lessingia vegetationmanagement.

continual *GGNRA

PT

7 per yr

(210)

7 7 7 

1 A 1.2.1 Establish protection of Daly City

population of San Francisco lessingiaunder existing ownership.

2 years

 or less

*USFWS, CDFG

DC

1 1 0 0 

1 A 1.2.2 Acquire Daly City parcels within

proposed San Francisco lessingiareserve.

2 years

or less

*USFWS,

CDFG,

PNP, DC, OWN

150 0 150 0 

1 A 2.1.1 Prepare plans for habitat restoration

and augmentation or reintroduction ofSan Francisco lessingia populations.

 4 years

or longer

(TBD)

*USFWS, CDFG

GGNRA, PT,

PNP, PC

25 10 5 5 

1 A 2.1.2 Do a genetics management plan for

San Francisco lessingia.

6 months *USFWS 6 6 0 0 

1 A 2.2.1 Expand population and restore habitat

above Lobos Dunes site, north andeast of Lobos Creek to BatteryCaulfield Road.

 10 years *GGNRA

USFWS

*PT, CDFG

75 0 0 48 

1 A 2.2.2 Expand populations and restore

habitat from Lobos Dunes to BakerBeach Dunes.

 5 years *GGNRA

USFWS

*PT, CDFG

40 0 0 20 

1 A.3.1 Prepare information presentations for

public outreach (San Franciscolessingia).

5 + years *USFWS,

CDFG, *PT, SF,

*GGNRA, PNP

TBD – – – 

208
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

1 A 3.2 Conduct public outreach meetings

prior to major recovery actions for SanFrancisco lessingia.

5 + years *USFWS, CDFG

*GGNRA, *PT,

SF, PNP

TBD – – – 

1 A 3.3 Publicize and promote restoration sites

for San Francisco lessingia in advanceof modifications.

5 + years *USFWS,

CDFG, GGNRA,

PT, SF, PNP

c. 1 per yr

(5)

111

1 A 3.5 Actively support public volunteer

participation in monitoring and sitestewardship of San Francisco lessingiareserves.

TBD USFWS, CDFG, 

PNP, *GGNRA,

SF, *PT

c. 1 per yr

(30)

111

1 A 3.6 Coordinate all outreach in actions 3.1-

3.5 with partners. Coordinatemethods, phasing, and prioritization.

30 *USFWS,

*GGNRA, *PT,

*SF

c. 1 per yr

(30)

111

1 A 4.1 Do genetic crossing studies as needed. 2 years *USFWS 1 0.5 0.5 0 1 A 4.5 Experimentally investigate rotational

trail closure/temporary pedestrianexclosure as a management andconservation method for SanFrancisco lessingia.

5+ years *USFWS,

CDFG,

*GGNRA, UNIV

3.5 1 1 0.5 

1 A 6. Develop a participation plan with all

partners.

1 year *USFWS,

*GGNRA, *PT,

*SF

1 100

1 B 1.1 Control nonnative vegetation at the

Presidio World War II MemorialRaven’s manzanita site.

ongoing *GGNRA, *PT 0.5 per yr

(15)

0.5 0.5 0.5 

1 B 1.3 Manage native vegetation around War

Memorial Raven’s manzanita site.

continual *GGNRA, *PT c. 0.5 per

yr

(15)

0.5 0.5 0.5 

209
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

1 B 2.2 Control nonnative vegetation at

Presidio transplant sites of Raven’smanzanita.

continual *GGNRA, *PT 0.5 per yr

(15)

0.5 0.5 0.5 

1 B 5.6 Investigate practical specific

nonnative vegetation controltechniques at Raven’s manzanita sites.

 3 years *USFWS, CDFG

GGNRA, UNIV,

PT

6 222

Total estimated costs for Priority 1 actions 663.9 40.6 176.3 88.25 

2 A 1.1.1.3.2 Evaluate West Washington Housing

for inclusion in Presidio RecoveryUnit.

1 year *USFWS, *PT,

GGNRA

0.6 0.6

2 A 1.1.2 Monitor all Presidio populations of

San Francisco lessingia.

continual *GGNRA, *PT TBD

c. 3 per yr

(90)

3 3 3 

2 A 1.2.3 Establish a land manager and

endowment fund for management ofthe Daly City San Francisco lessingiareserve.

2 years

or less

*USFWS,

CDFG,

DC, PNP

75 0 0 75 

2 A 1.2.4 Prepare and implement a long-term

management and restoration plan forDaly City San Francisco lessingiareserve.

3 years *USFWS, CDFG

DC, PNP, PC

20 0 10 5 

210
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

2 A 2.2.3 Expand San Francisco lessingia

populations and restore habitat aboveLobos Dune Reserve through WherryHousing Area, Presidio.

30 years *PT, *GGNRA,

USFWS, CDFG

550 0 0 0 

2 A 2.2.4 Expand San Francisco lessingia

populations and restore habitat behindPublic Health Services Hospital,Presidio.

4 years *PT, GGNRA,

USFWS, CDFG

75 1 4 65 

2 A 2.2.5 Expand San Francisco lessingia

populations and restore habitat withinthe Daly City Reserve.

3 years *USFWS,

CDFG, DC, PC

(20) 

2 A 2.2.6 Restore habitat and reintroduce San

Francisco lessingia to dunes at FortFunston reserve for San Franciscolessingia.

5 years *GGNRA,

USFWS, CDFG

205 5 50 50 

2 A 2.3 Reintroduce beach layia to Fort

Funston dunes.

4 years *GGNRA,

USFWS, CDFG

3 1 1 0.5 

211
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

2 A 2.4.1 Acquire Hawk Hill dune remnant. 1 year *USFWS,

CDFG, SF, PNP,

OWN

150 150 0 0 

2 A 2.4.2 Determine ownership and acquire

sand slopes near Sutro Heights.

2 years *SF, OWN,

GGNRA,

USFWS, CDFG

150 5 145 0 

2 A 2.4.3 Control or eradicate invasive

nonnative vegetation within satellitereserves.

continual *GGNRA, PNP,

USFWS, *SF

150 5 5 5 

2 A 2.4.4 Establish San Francisco lessingia in

satellite reserves.

5 years *USFWS,

CDFG, GGNRA,

SF, PNP

30 5 10 5 

2 A 3.4 Publicize effects of San Francisco

lessingia projects on scenic views andrecreational use of GGNRA/Presidio.

5 + years *USFWS,

CDFG, *PT, SF,

*GGNRA, PNP

c. 3 per yr

(15)

333

2 A 4.3 Study rates and modes of

recolonization by invasive nonnativeplants and their control.

5+ years *USFWS,

CDFG, GGNRA,

UNIV

5 2 1 1 

2 A 4.6 Conduct field experiments on

restoring and managing dunetopography conservation of SanFrancisco lessingia.

5+ years *USFWS,

CDFG, GGNRA,

UNIV

7.5 5 1 0.5 

2 A 4.8 Study effects of substrate conditions

on growth and reproduction of SanFrancisco lessingia.

5 years *USFWS,

CDFG, GGNRA,

UNIV

7 221

212
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

2 A 5.1 Establish cultivated populations of

San Francisco lessingia at botanicalgardens.

ongoing *USFWS,

CDFG, UNIV,

PNP 

1 0.5 0.5 0 

2 B 1.2 Monitor growth, reproduction, and

clone size of Raven’s manzanita at theWorld War II Memorial manzanitasite.

continual *GGNRA, *PT ca. 0.5 per

yr

(15)

0.5 0.5 0.5 

2 B 2.1 Monitor growth and clone size of

established Raven’s manzanitatransplants in the Presidio.

continual *GGNRA, *PT up to

0.1 per yr

(3)

0.1 0.1 0.1 

2 B 2.3 Survey and select additional

serpentine outcrop sites within thePresidio for reintroduction of Raven’smanzanita.

2 years USFWS, CDFG,

*GGNRA, *PT

3 1.5 1.5 0 

2 B 2.4 Prepare sites selected for

transplantation of Raven’s manzanitain the Presidio.

TBD *USFWS,

CDFG, GGNRA,

PT

TBD – – – 

2 B 2.5 Transplant replicated clones of

Raven’s manzanita to selected sites inthe Presidio.

3 years *USFWS,

CDFG, GGNRA,

PT

3 1 1 1 

2 B 2.6 Monitor transplanted Presidio clones. 7 years USFWS, CDFG,

*GGNRA, *PT

7 0 0 0 

2 B 2.7 Identify and protect known and

potential serpentine outcrop sites inthe Presidio. 

3 years *GGNRA, *PT,

PC

5 131

2 B 3.1 Evaluate and select candidate reserves

for reintroduced Raven’s manzanitaoutside the Presidio in San Francisco.

1 year *USFWS,

CDFG, PNP, SF,

OWN

1 100

213
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

2 B 3.2 Acquire selected Raven’s manzanita

reserves outside the Presidio in SanFrancisco.

TBD (3

years?)

*USFWS,

CDFG, PNP,

SF, OWN

TBD – – – 

2 B 3.3 Prepare restoration and reintroduction

plans for Raven’s manzanita reservesoutside the Presidio in San Francisco.

2 years *USFWS,

CDFG, SF, PNP,

PC, UNIV

5 023

2 B 3.4 Propagate native manzanita stock

from seed.

5 years *USFWS,

CDFG, UNIV,

PNP

10 2 2 2 

2 B 3.5 Prepare new San Francisco sites for

reintroduction of native manzanitas.

2 years *USFWS,

CDFG, PNP,

PC, SF

20 0 0 0 

2 B 3.6 Transplant and establish propagated

native founder manzanita populations.

3 years *USFWS,

CDFG, PNP,

PC, SF

1 0 0 0 

2 B 4.1 Promote native manzanita recovery

through information and outreachefforts.

5+ years *GGNRA,

USFWS, CDFG, 

PNP, SF, UNIV

5 1 1 1 

2 B 4.2 Promote native manzanita recovery at

the local community level.

5+ years *GGNRA,

USFWS, CDFG, 

PNP, SF, UNIV

8 1 1 2 

2 B 5.1 Search for “Raven’s-like” Tamalpais

manzanitas (Arctostaphylos montana).

3(+) years *USFWS,

CDFG, UNIV

1 0.2 0.2 0.2 

2 B 5.2 Investigate seed production and

propagation of Raven’s manzanita.

4 years *UNIV,

USFWS, CDFG

6 2 2 1 

214
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

2 B 5.3 Investigate taxonomic status and

relationships of Raven’s manzanita.

3 years *UNIV,

USFWS, CDFG

8 431

2 B 5.4 Investigate substrate and plant

interactions with Raven’s manzanita.

5 years *UNIV,

USFWS, CDFG

9 322

2 B 5.5 Investigate reproductive ecology of

Raven’s manzanita.

3 years *UNIV,

USFWS, CDFG

5 221

2 B 5.7 Assemble a scientific review panel to

evaluate and recommend a geneticmanagement plan for Raven’smanzanita.

1 year *USFWS,

CDFG, UNIV,

PC, GGNRA,

*PT

2 200

Total estimated costs for priority 2 actions 1,651.1 210.4 256.8 229.8 

3 A 4.2 Study variation in ecological and

genetic characteristics of SanFrancisco lessingia.

2 years *USFWS,

CDFG, GGNRA,

UNIV, PC, PT

5 0 2 3 

3 A 4.4 Study the pollination ecology of San

Francisco lessingia.

2 years *USFWS,

CDFG, GGNRA,

UNIV, PT

2 110

3 A 4.7 Study seed dispersal and seed bank

ecology of San Francisco lessingia.

5+ years *USFWS,

CDFG, GGNRA,

UNIV, PT

5 0 2 2 

3 A 4.9 Clarify taxonomic relationships

between San Francisco lessingia andallied species.

4 years *USFWS,

CDFG,

GGNRA,

UNIV, DC,

OWN, PT

5 211

3 A 5.2 Maintain stored seed bank to ensure

survival of San Francisco lessingia. 

ongoing *UNIV, PNP,

USFWS

3 0.1 0.1 0.1 

215
Action

Priority 

ActionNumber Action Description †

Action

Duration

Responsible

Parties

Cost Estimate (in $10,000 units) by f

Total ‡ 1 2 3 

3 B 2.8 Reintroduce associated serpentine

plant species to serpentine outcropsites in the Presidio.

ongoing *GGNRA, *PT,

SF, PNP

TBD – – – 

3 B 3.7 Monitor transplants of Raven’s

manzanita in San Francisco outsidethe Presidio.

ongoing *UNIV, PC,

PNP, USFWS,

CDFG

TBD – – – 

3 B 4.3 Promote native manzanita recovery

through stewardship.

continual *GGNRA, *PT

USFWS, CDFG, 

PNP, SF,

TBD – – – 

3 B 4.4 Establish public demonstration

gardens displaying cultivated Raven’smanzanita clones.

5 years *GGNRA, PNP,

USFWS, CDFG,

SF, *PT

1.25 0.25 0.25 0.25 

3 B 6.1 Collect and store seed of Raven’s

manzanita.

ongoing *GGNRA, *PT

UNIV, PNP

3 0.1 0.1 0.1 

3 B 6.2 Cultivate Raven’s manzanita in

botanical gardens.

ongoing *PNP, UNIV,

USFWS, CDFG

1 1 0 0 

3 B 6.3 Promote and authorize appropriate

informational and outreach use ofRaven’s manzanita.

ongoing *USFWS,

CDFG, PNP, SF

3 0.1 0.1 0.1 

Total estimated costs for priority 3 actions 28.25 4.55 6.55 6.55 Total estimated costs 2,343.25 255.55 439.65 324.60 

† Action Description: Please see Stepdown Narrative (Chapter V) for a full list of species included in eac

216
217

‡ For purposes of estimating total cost of implementing the recovery plan (see Executive Summary), conactions or actions of undetermined duration with costs given on a per year basis are multiplied by 30 yearestimate of time to delisting of San Francisco lessingia and downlisting of Raven’s manzanita. Actions oduration with costs given on a per year basis are multiplied by the duration of the action.
218

VII. REFERENCES

A. Literature Cited

Abrams, L. 1944. Illustrated flora of the Pacific states. Volume II. Stanford

University Press. 635 pp.

________. 1951. Illustrated flora of the Pacific states. Volume III. Stanford

University Press. 866 pp.

Albert, M. E., C. M. D’Antonio, and K. A. Schierenbeck. 1997. Hybridization and

introgression in Carpobrotus spp. (Aizoaceae) in California. I. Morphological

evidence. American Journal of Botany 84:896-904.

Atwater, B. F., S. G. Conard, J. N. Dowden, C. H. Hedel, R. L. MacDonald, and W.

Savage. 1979. History, landforms and vegetation of the estuary’s tidal

marshes, in San Francisco Bay: the urbanized estuary: investigations into the

natural history of San Francisco Bay and Delta with reference to the influence

of man. 58th annual meeting of the American Association of the Advancement

of Science, Pacific Division, ed T. J. Conomos. Pacific Division, AAAS. 493

pp.

Barrett, C. H., and J. R. Kohn. 1991. Genetic and evolutionary consequences of small

population size in plants: implications for conservation. Pp. 3-32 in Genetics

and conservation of rare plants, eds. D. A. Falk and K. E. Holsinger. Oxford

University Press, New York. 281 pp.

Beardsmore, J. A. 1983. Extinction, survival and genetic variation. Pp. 125-151 in 

Genetics and conservation, eds. C. M. Schoenwald-Cox, S. M. Chambers, B.

MacBryde, and L. Thomas. Benjamin-Cummings, Menlo Park, California.

Behr, H. H. 1888. Flora of the vicinity of San Francisco. San Francisco, California.

________. 1892. Botanical Reminiscences. Zoe 2:2-6.
219

Best, C., J. T. Howell, W. and I. Knight, and M. Wells. 1996. A flora of Sonoma

County. California Native Plant Society. 347 pp.

Bode, K. A. 1998. Report on Lessingia germanorum monitoring protocols. Report to

Presidio Natural Resources, Golden Gate National Recreation Area, National

Park Service, Fort Mason, San Francisco, California.

Bolander, H. N. 1863. Enumeration of shrubs and trees growing in the vicinity of the

mouth of San Francisco Bay. California Academy of Sciences Proceedings

2:78-84.

Bonilla, M. G. 1965. Geologic map of the San Francisco South quadrangle,

California: U.S. Geological Survey open-file map, scale 1:20,000.

Brandegee, K. 1892. Catalogue of the flowering plants and ferns growing

spontaneously in the city of San Francisco. Zoe 2:334-383.

Briggs, F. N. and P. F. Knowles. 1967. Introduction to plant breeding. Reinhold

Pub. Co., New York. 426 pp.

Burdon, J. J. 1987. Diseases and plant population biology. Cambridge University

Press, Cambridge and New York. 208 pp.

CALFLORA 1999. on-line. California floristic database, a collaborative project of

the U.S. Department of Agriculture Forest Service; University of California,

Berkeley, Digital Library Project; the Bioinformatics Working Group of Texas

A&M University; the University of California, Davis, Information Center for

the Environment; the Santa Barbara Botanical Garden; and others. 

http://galaxy.cs.berkeley.edu/calflora.

California Department of Fish and Game. 1992. Annual report on the status of

California State listed threatened and endangered animals and plants. 

California Department of Fish and Game, Sacramento, California. 

Carroll, R., C. Augsperger, A. Dobson, J. Franklin, G. Orians, W. Reid, R. Tracy, D.

Wilcove, and J. Wilson. 1996. Strengthening the use of science in achieving
220

the goals of the Endangered Species Act: An assessment by the Ecological

Society of America. Ecological Applications: 6:1-11.

Carter, R. W. G. 1988. Coastal environments. Academic Press, Harcourt Brace

Jovanovich, Publishers. London, New York. 617 pp.

Center for Plant Conservation. 1991. Genetic sampling guidelines for conservation

collections. Pp. 225-239 in Genetics and conservation of rare plants, eds. D.

A. Falk and K. E. Holsinger. Oxford University Press, New York. 281 pp.

Chuang, T. I., and L. R. Heckard. 1993. Castilleja. Pp.1016-1024 in The Jepson

manual: higher plants of California, ed. J. C. Hickman. University of

California Press. 

1,400 pp.

Clary, R. H. 1980. The making of Golden Gate Park. California Living Books, San

Francisco Examiner Division of the Hearst Corporation, San Francisco.

Cooper, J. G. 1875. The influence of climate and topography on our trees. 

Proceedings of the California Academy of Sciences 5(3):285-291.

Cooper, W. S. 1936. The strand and dune flora of the Pacific coast of North America: 

a geographic study. Pp. 147-187 in Essays in geobotany in honor of William

Albert Setchell, ed. T. H. Goodspeed. University of California Press. 319 pp.

________. 1958. Coastal sand dunes of Oregon and Washington. Geological Society

of America Memoir 72. 169 pp.

________. 1967. Coastal dunes of California. Geological Society of America

Memoir 104. 131 pp.

Crow, J. F. 1988. The importance of recombination. Pp. 56-73 in The evolution of

sex: an examination of current ideas, ed. R. E. Michod and B. E. Levin. 

Sinauer Associates, Inc, Sunderland, Massachusetts. 342 pp.
221

Davies, J. L. 1980. Geographical variation in coastal development, 2nd ed. 

Longman, New York, 212 pp.

Day, A. G. 1993. Gilia. Pp. 828-836 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Dewey, S. A. 1995. Noxious Weeds, A Biological Wildfire, Applying Fundamentals

of Wildfire Management to Improve Noxious Weed Control. AG 500 (10M/8-

95). Utah State University Extension; Logan, Utah. 6pp.

DuRietz, G. E. 1930. The fundamental units of biological taxa. Svensk. Bot. Tidskr. 

24:333-428.

Ellstrand, N. C. 1992. Gene flow by pollen: implications for plant conservation

genetics. Oikos 3:77-86.

Ellstrand, N. C., J. M. Lee, J. E. Keeley, and S. C. Keeley. 1987. Ecological isolation

and introgression: biochemical confirmation of introgression in an

Arctostaphylos population. Acta Oecologica/ Oecologia Plantarum 8:299-308.

Elzinga, C. L., Salzer, D. W., and J. W. Willoughby. 1998. Measuring and

monitoring plant populations. BLM Technical Reference 1730-1. 477 pp.

Ertter, B. 1993. Horkelia. Pp. 953-957 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Faegri, K., and L. van der Pijl. 1979. The principles of pollination ecology, 3rd

revised edition. Pergamon Press; Oxford, New York. 244 pp.

Falk, D. A. 1992. From conservation biology to conservation practice: strategies for

protecting plant diversity. Pp. 373-432 in Conservation biology: the theory

and practice of conservation, preservation, and management, eds. Peggy L.

Fiedler and Subodh K. Jain. Chapman and Hall, New York. 507 pp. 

Falk, D. A., C. I. Millar, and M. Olwell. 1996. Guidelines for developing a rare plant

reintroduction plan. Pp. 453-491.
222

Fimrite, Peter. 2001. Oak-killing disease spreads to buckeyes; latest victims, some 

discovered in Bay Area, are the 10th species stricken by fungus-like organism. 

San Francisco Chronicle, Monday, July 2, 2001. San Francisco, California.

Frankel, O. H., and M. Soulé. 1981. Conservation and evolution. Cambridge

University Press; Cambridge, New York. 327 pp.

Fraser, G. S., S. W. Bennett, G. A. Olyphant, N. J. Bauch, V. Ferguson, C. A.

Gellasch, C. L. Millard, B. Mueller, P. J. O’Malley, J. H. Way, and M. C.

Woodfield. 1998. Windflow circulation patterns in a coastal dune blowout,

south coast of Lake Michigan. Journal of Coastal Research 14:451-460.

Gallagher, K. G., K. A. Schierenbeck, and C. M. D’Antonio. 1997. Hybridization and

introgression in Carpobrotus spp. (Aizoaceae) in California. II. Allozyme

evidence. American Journal of Botany 84:905-911.

Gares, P. A., and K. F. Nordstrom. 1990. Topographic changes in a formerly

stabilized dune system in a coastal park. Pp. 159-170 in Proceedings of the

symposium on coastal sand dunes, September 12-14, 1990, Guelph, Ontario,

ed. R. Davidson-Arnott. Coastal Zone Engineering, Institute for Mechanical

Engineering, National Research Council Canada, Ottawa, Canada.

Ghiselin, M. T. 1988. The evolution of sex: a history of competing points of view. 

Pp. 7-23 in The evolution of sex: an examination of current ideas, ed. R. E.

Michod and B. E. Levin. Sinauer Associates, Inc, Sunderland, Massachusetts. 

342 pp.

Golden Gate National Recreation Area. 1997. Of dogs and parks: striking a balance.

Page 2, ParkLands, the quarterly magazine of the National Parks at the Golden

Gate, Dec-Feb, Winter 1996-1997 issue.

Golden Gate National Recreation Area and Presidio Trust. 2000. Presidio Vegetation 

Management Plan. National Park Service and The Presidio Trust, Building

204, Fort Mason, San Francisco, California.
223

Gottlieb, L. D. 1968. Hybridization between Arctostaphylos viscida and A. canescens

in Oregon. Brittonia 20:83-93.

________. 1973. Enzyme differentiation and phylogeny in Clarkia franciscana, C.

rubicunda, and C. amoena. Evolution 27:205-214.

Gottlieb, L. D., and S. W. Edwards. 1992. Electrophoretic test of the genetic

independence of a newly discovered population of Clarkia franciscana. 

Madroño 39:1-7.

Greene, E. L. 1894. Manual of the botany of the region of San Francisco Bay.

Cubery and Company, San Francisco. 342 pp.

Guerrant, E. O. 1992. Genetic and demographic considerations in the sampling and

reintroduction of rare plants. Pages 321-344 in Conservation biology: the

theory and practice of conservation, preservation, and management, eds.

Peggy L. Fiedler and Subodh K. Jain. Chapman and Hall, New York. 507 pp.

________. 1996. Designing populations: demographic, genetic, and horticultural

dimensions. Pages 171-208 in Restoring diversity: strategies for

reintroduction of endangered plants, eds. Donald A. Falk, Constance I. Millar,

Margaret Olwell. Island Press; Washington D.C. 505 pp.

Heckard, L. R. 1993. Orobanchaceae (including Orobanche). Pp. 804-808 in The

Jepson manual: higher plants of California, ed. J. C. Hickman. University of

California Press. 1,400 pp.

Hickman, J. C. (ed.). 1993. The Jepson manual: higher plants of California. 

University of California Press. Berkeley, California. 1,400 pp. [note:

Hickman wrote some treatments, including Chorizanthe, pp. 856-860].

Holsinger, K. E., and L. D. Gottlieb. 1991. Conservation of rare and endangered

plants: principles and prospects. Pp. 195-208 in Genetics and conservation of

rare plants, eds. D. A. Falk and K. E. Holsinger. Oxford University Press,

New York. 281 pp.
224

Hoover, R. F. 1970. The vascular plants of San Luis Obispo County, California. 

University of California Press, Berkeley. 350 pp.

Howard, A. D. 1979. Geologic history of middle California. University of California

Press, Berkeley. 113 pp.

Howell, J. T. 1929. A systematic study of the genus Lessingia Cham. University of

California Publications in Botany, vol. 16. University of California Press,

Berkeley. 44 pp.

________. 1949. Marin flora. University of California Press, Berkeley. 319 pp.

Howell, J. T., P. H. Raven, and P. R. Rubtzoff. 1958. A flora of San Francisco,

California. The Wasmann Journal of Biology 16:1-157.

Huenneke, L. F. 1991. Ecological implications of genetic variation in plant

populations. Pp. 31-44 in Genetics and conservation of rare plants, eds. D. A.

Falk and K. E. Holsinger. Oxford University Press, New York. 281 pp.

Isely, D. 1993. Trifolium. Pp. 646-654 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Jepson, W. L. 1911. A Flora of Western Middle California. Cunningham, Curtiss

and Welch, San Francisco. 515 pp. 2nd ed. (1st ed.1901).

________. 1925. Manual of the Flowering Plants of California. Associated Students

Store, University of California, Berkeley. 24, 24a-24h, 25-1238 pp.

Jones and Stokes. 1996. Environmental Assessment: Crissy Field Plan, Golden Gate

National Recreation Area. (JSA 995-127) Sacramento, California. Prepared

for National Park Service, Golden Gate National Recreation Area, Fort Mason,

San Francisco, California.

Jungerius, P. D., A. J. T. Verheggen, and A. J. Wiggers. 1981. The development of

blowouts in ‘de Blink,’ a coastal dune area near Noordwijkerhout, the

Netherlands. Earth Surface Processes and Landforms 6:375-396.
225

Kaufeldt, C. E. 1954. Observations of San Francisco’s last major sand dune, with

notes on its flora and insect fauna. The Wasmann Journal of Biology 12:293-

336.

Keeley, J. E. 1976. Morphological evidence of hybridization between Arctostaphylos

glauca and A. pungens (Ericaceae). Madroño 23:427-433.

________. 1987. Role of fire in seed germination of woody taxa in California

chaparral. Ecology 68:434-443.

Keil, D. J., and C. E. Turner. 1993. Cirsium, pp. 232-239 in The Jepson manual: 

higher plants of California, ed. J. C. Hickman. University of California Press. 

1,400 pp.

Kelly, V. R., and V. T. Parker. 1990. Seed bank survival and dynamics in sprouting

and nonsprouting Arctostaphylos species]. American Midland Naturalist

124:114-123. 

________. 1991. Percentage seed set, sprouting habit and ploidy level in

Arctostaphylos (Ericaceae). Madroño 38:227-232.

Knapp, E. E., and P. G. Connors. 1999. Genetic consequences of a single-founder

population bottleneck in Trifolium amoenum (Fabaceae). American Journal of

Botany 

86:124-130.

Knapp, E. E., and K. J. Rice. 1994. Starting from seed: genetic issues in using native

grasses for restoration. Restoration and Management Notes 12:40-445.

Kruckeberg, A. R. 1954. The ecology of serpentine soils. III. Plant species in

relation to serpentine soils. Ecology 35:267-274.

________. 1977. Manzanita (Arctostaphylos) hybrids in the Pacific Northwest: 

effects of human and natural disturbance. Systematic Botany 2:233-250.
226

________. 1984. California serpentines: flora, vegetation, geology, soils, and

management problems. University of California Press, Berkeley. 180 pp.

Kutner, L. S., and L. E. Morse. 1996. Reintroduction in a changing climate. Pp. 23-

48 in Restoring diversity: strategies for reintroduction of endangered plants,

eds. Donald A. Falk, Constance I. Millar, Margaret Olwell. Island Press;

Washington D.C. 505 pp.

Kyhos, D. W., and P. H. Raven. 1982. Miscellaneous chromosome numbers in

Asteraceae. Madroño 29:62.

Lane, M. Grindelia. Pp. 271-274 in The Jepson manual: higher plants of California,

ed. J. C. Hickman. University of California Press. 1,400 pp.

Lane, M. Lessingia. Pp. 304-306 in The Jepson manual: higher plants of California,

ed. J. C. Hickman. University of California Press. 1,400 pp.

Leatherman, S. P, ed. 1979. Barrier islands from the Gulf of St. Lawrence to the Gulf

of Mexico. Academic Press, New York. 325 pp.

Lenz, L. 1956. Native plants for California gardens. Rancho Santa Ana Botanic

Garden, Day Printing Corporation, Pomona, California. 166 pp.

Lesica, P., and F. Allendorf. 1999. Ecological genetics and the restoration of plant 

communities: mix or match? Restoration Ecology 7:42-50.

Lesica, P., and S. Cooper. 1999. Succession and disturbance in sandhills vegetation: 

constructing models for managing biological diversity. Conservation Biology 

13:293-303.

Levin, D. A., J. Francisco-Ortega, and R. K. Jansen. 1996. Hybridization and the

extinction of rare plant species. Conservation Biology 10:10-16.

Lewis, H. 1993. Clarkia. Pp. 786-793 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.
227

Lewis, H., and P. H. Raven. 1958. Rapid evolution in Clarkia. Evolution 12:319-

336.

Mace, G. M. and R. Lande. 1991. Assessing extinction threats: Toward a

reevaluation of IUCN threatened species categories. Conservation Biology

5:148-157.

Markos, S., and B.G. Baldwin. 2002. Structure, molecular evolution, and

phylogenetic utility of the 5' region of the external transcribed spacer of 18S26S rDNA in Lessingia (Compositae, Astereae). Molecular Phylogenetics and

Evolution 23(2):214-228.

Markos, S., L. C. Hileman, M. C. Vasey, and V. T. Parker. 1999. Phylogeny of the

Arctostaphylos hookeri complex (Ericaceae) based on nrDNA data. Madroño

45:187-199.

Matthews, M. A. 1997. An illustrated field key to the flowering plants of Monterey

County and ferns, fern allies, and conifers. California Native Plant Society

Publications, Sacramento. 401 pp.

McCarten, N. F. 1986. A study of the ecological aspects related to the reintroduction

of Arctostaphylos hookeri ssp. ravenii. Unpublished report submitted to

California Department of Fish and Game, Endangered Plant Project.

McClintock, E. 1993. Tanacetum. P. 350 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

McClintock, E., P. Reeberg, and W. Knight. 1990. A Flora of the San Bruno

Mountains, San Mateo County, California [rev. and updated ed.]. California

Native Plant Society Special Publication number 8. Sacramento, California.

223 pp.

McLaren, J. 1924. Gardening in California: landscape and flower. A. M. Robertson 

Publishers, San Francisco. 3rd ed. 395 pp.
228

McMahon, L. R. 1989. Rarest U.S. plants are literally one of a kind. Plant

Conservation 4:6.

McMinn, H. E. 1939. An illustrated manual of California shrubs. J. W. Stacey, Inc.,

San Francisco. 689 pp. [reprints, 1951 onward, by Univ. of California Press,

Berkeley]. 

Messick, T. C. 1993. Plagiobothrys. Pp. 386-390 in The Jepson manual: higher

plants of California, ed. J. C. Hickman. University of California Press. 1,400

pp.

Millar, C. I., and W. Libby. 1991. Strategies for conserving clinal, ecotypic, and

disjunct population diversity in widespread species. Pages 149-170 in

Genetics and conservation of rare plants, eds. D. A. Falk and K. E. Holsinger. 

Oxford University Press, New York. 281 pp.

Miller, R., D. Loeb, J. Eaton, R. Sullivan, and C. Clarke. 1997. The plover’s best

friend. Terrain, Northern California’s Environmental Magazine, April 1997.

Mollette, Y. 1998. Sand dune coastal scrub seed bank analysis and implications of

patch dynamics. M. A. thesis, San Francisco State University.

Morefield, J. D. 1993. Hesperevax. P. 284 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Munz, P. A. 1959. A California flora. University of California Press, Berkeley. 

1,681 pp.

National Park Service. 1994. Final general management plan amendment

environmental impact statement, Presidio of San Francisco, Golden Gate

National Recreation Area, California. Golden Gate National Recreation Area,

Fort Mason, San Francisco, California.

________. 2003. Presidio of San Francisco, Golden Gate National Recreation Area. 

Special Status Plant Species Monitoring Report, 2001-2002. Internal report
229

prepared by Tim Doherty for Presidio Natural Resources, GGNRA, NPS, San

Francisco, California. 55 pp.

National Park Service and Presidio Trust. 2001. Presidio of San Francisco,

Vegetation Management Plan and Environmental Assessment. Golden Gate

National Recreation Area and Presidio Trust, San Francisco, California. 

109pp. + appendices.

National Research Council. 1995. Science and the Endangered Species Act. National

Academy Press; Washington, DC. 271pp.

Nesom, G. L. 1993. Erigeron. Pp. 253-261 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Nunney, L., and K. A. Campbell. 1993. Assessing minimum viable population size: 

demography meets population genetics. Trends in Ecology and Evolution

8:234-238.

Olson, J. S. 1958. Lake Michigan dune development. I. Wind velocity profiles. 

Journal of Geology 66:254-263.

Patterson, R. W. 1993. Linanthus. Pp. 840-844 in The Jepson manual: higher plants

of California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Pennell, F. W. 1951. Castilleja, in Illustrated Flora of the Pacific States, vol. 3. 

L. Abrams. Stanford University Press, Stanford, Calif.

Pogge, C. 1998. Life History and Ecology of Lessingia germanorum. M.A. thesis,

San Francisco State University.

Powell J. A., and C. L. Hogue. 1979. California insects. University of California

Press, Berkeley. 388 pp.

Presidio Trust. 2001. Presidio Trust Implementation Plan Environmental Impact 

Statement. The Presidio Trust, 34 Graham Street, San Francisco, California.
230

Presidio Trust. 2002. Presidio Trust Management Plan. The Presidio Trust, 34

Graham Street, San Francisco, California. 136 pp. + appendices.

Primack, R. B. 1996. Lessons from ecological theory: dispersal, establishment, and

population structure. Pages 209-234 in Restoring diversity: strategies for

reintroduction of endangered plants, eds. Donald A. Falk, Constance I. Millar,

Margaret Olwell. Island Press; Washington D.C. 505 pp.

Proctor, M., P. Yeo, and A. Lack. 1996. The Natural History of Pollination. Timber

Press, Portland, Oregon. 479 pp.

Ramaley, F. 1918. Notes on dune vegetation at San Francisco. Plant World 21:191-

201.

Ranwell, D. S. 1972. Ecology of salt marshes and sand dunes. Chapman and Hall,

London. 258 pp.

Raven, P. H. 1952. Plant notes from San Francisco, California. Leaflets of Western

Botany 6:208-211.

Raven, P. H. and D. Axelrod. 1978. Origin and Relationships of the California Flora. 

University of California Publications in Botany, Vol. 72. Univ. of California

Press, Berkeley. 134 pp.

Reinartz, J. A. 1995. Planting state-listed endangered and threatened plants. 

Conservation Biology 9:771-781.

Rieseberg, L. H. 1991. Hybridization in rare plants: insights from case studies in

Cercocarpus and Helianthus. Pp.171-181 in Genetics and conservation of

rare plants, eds. D. A. Falk and K. E. Holsinger. Oxford University Press,

New York. 281 pp.

Robichaux, R., H., E. A. Friar, and D. W. Mount. 1997. Molecular and genetic

consequences of a population bottleneck associated with reintroduction of the

Mauna Kea Silversword (Argyroxiphium sandwichense ssp. sandwichense

[Asteraceae]). Conservation Biology 11:1140-1146.
231

Roof, J. 1971. Notice of Clarkia franciscana sowings. The Four Seasons 4:17.

________. 1972. Summer wildflowers: the Clarkias. The Four Seasons 4:2-6.

________. 1976. A fresh approach to the genus Arctostaphylos in California. 

The Four Seasons 5:20-24.

________. 1978. Studies in Arctostaphylos (Ericaceae). The Four Seasons 5:3-15.

________. 1980. California’s Arctostaphylos uva-ursi alliance. The Changing

Seasons 1(2):19-26.

Schemske, D. W., B. C. Husband, M. H. Ruckelshaus, C. Goodwillie, I. M. Parker,

and J. G. Bishop. 1994. Evaluating approaches to the conservation of rare and

endangered plants. Ecology 75:584-606.

Schierenbeck, K. A., G. L. Stebbins, and R. W. Patterson. 1992. Morphological and

cytological evidence for polyphyletic allopolyploidy in Arctostaphylos

mewukka (Ericaceae). Plant Systematics and Evolution 179:187-205.

Schlocker, J. 1974. Geology of the San Francisco North Quadrangle, California. 

U.S. Geological Survey Professional Paper 782, United States Government

Printing Office, Washington, D.C.

Scott, J. M., T. H, Tear, and L. S. Mills. 1995. Socioeconomics and the recovery of

endangered species: Biological assessment in a political world. Conservation

Biology 9:214-216.

Shields, W. M. 1988. Sex and adaptation. Pages 253-286 in The evolution of sex: 

an examination of current ideas, ed. R. E. Michod and B. E. Levin. Sinauer

Associates, Inc, Sunderland, Massachusetts. 342 pp.

Skinner, M. W., and B. M. Pavlik. 1994. California Native Plant Society’s inventory

of rare and endangered vascular plants of California, 5th edition. California

Native Plant Society Publications, Sacramento, California. 338 pp. 
232

Smith, G. L. 1986. Taxonomic and nomenclatural notes on Tanacetum camphoratum

and Tanacetum douglasii (Asteraceae). The Wassmann Journal of Biology

44:130-134.

Smith, G. L., and C. R. Wheeler. 1990-1991. A Flora of the vascular plants of

Mendocino County. Reprinted from The Wasmann Journal of Biology vols.

48-49, nos. 1 and 2, spring and fall 1990-1991, pp. 1-387. University of San

Francisco, San Francisco, California.

Spellenberg, R. 1993a. Nyctaginaceae [including Abronia]. Pp. 768-774 in The

Jepson manual: higher plants of California, ed. J. C. Hickman. University of

California Press. 1,400 pp.

Spellenberg, R. 1993b. Astragalus. Pp. 583-605 in The Jepson manual: higher

plants of California, ed. J. C. Hickman. University of California Press. 1,400

pp.

Spence, W. L. 1964. A Biosystematic Study of the Genus Lessingia Cham.

(Compositae). Ph.D. dissertation, University of California.

Stebbins, G. L. 1950. Variation and Evolution in Plants. Columbia University Press,

New York. 643 pp.

Taylor, D., and D. H. Wilken. 1993. Atriplex. Pp. 501-505 in The Jepson manual: 

higher plants of California, ed. J. C. Hickman. University of California Press. 

1,400 pp.

Tear, T. H, J. M. Scott, P. H. Hayward, and B. Griffith. 1993. Status and prospects

for success of the Endangered Species Act: A look at recovery plans. Science

262:976-977.

Thomas, J. H. 1961. Flora of the Santa Cruz Mountains of California, a manual of

the vascular plants. Stanford University Press, Stanford, Calif. 434 pp. 
233

Thompson, K. 1992. The functional ecology of seed banks. Pp. 231-258 in Seeds: 

the ecology of regeneration in plant communities, ed. M. Fenner. C.A.B. 

International, Wallingford, Oxon, UK. 373 pp.

Tillett, S. S. 1967. The maritime species of Abronia (Nyctaginaceae). Brittonia

19:299-327.

U.S. Fish and Wildlife Service. 1979. Determination that Arctostaphylos hookeri ssp.

ravenii is an endangered species. Federal Register 44(209):61910-61911. 

Friday, October 26, 1979.

________. 1980. Endangered and threatened wildlife and plants: review of plant taxa

for listing as endangered or threatened. Federal Register 45(242):82480-

82569. December 15, 1980.

________. 1983a. Endangered and threatened species: Listing and recovery priority

guidelines. Federal Register 48:43098-43105. September 21, 1983.

________. 1983b. Endangered and threatened species: Listing and recovery priority

guidelines correction. Federal Register 48:51982. November 15, 1983.

________. 1984. Raven’s manzanita recovery plan. U.S. Fish and Wildlife Service,

Portland, Oregon.

________. 1993. Endangered and threatened wildlife and plants; endangered status

for two plants, Arenaria paludicola (marsh sandwort) and Rorippa gambellii

(sic) (Gambel’s watercress). Federal Register 58(147):41378-41384. Tuesday,

August 3, 1993.

________. 1994a. Endangered and Threatened Wildlife and Plants; Endangered and

threatened status for five plants and the Morro shoulderband snail from

western San Luis Obispo County, California. Federal Register 59(240):64613-

64623. Thursday, December 15, 1994.

________. 1994b. Endangered and Threatened Wildlife and Plants; Proposed

Endangered Status for the Plant Lessingia germanorum (San Francisco
234

Lessingia) and Threatened Status for the Plant Arctostaphylos imbricata (San

Bruno Mountain manzanita) from California. Federal Register 59(191):50550-

50557. Tuesday, October 4, 1994.

________. 1997. Endangered and threatened wildlife and plants; endangered status

for the plant Lessingia Germanorum [sic] (San Francisco Lessingia) from

California. Federal Register 62 (118):33368-33374. Thursday, June 19, 1997.

________. 1998a. Recovery plan for serpentine soil species of the San Francisco Bay

area. Portland, Oregon. 330 + pp.

________. 1998b. Recovery plan for seven coastal plant species and Myrtle’s

silverspot butterfly. Portland, Oregon. 141 + pp.

________. 1998c. Recovery Plan for the Marsh Sandwort and Gambel’s Watercress. 

Portland, Oregon. 48 + pp.

________. 1998d. Recovery plan for Kokia cookei. Portland, Oregon. 55 pp. +

appendices.

________. 2000. Policy Regarding Controlled Propagation of Species Listed Under

the Endangered Species Act. Federal Register 65(183):56916-56922.

________. 2001. Availability of Draft Recovery Plan for Coastal Plants of the

Northern San Francisco Peninsula for Review and Comment. Federal Register

66(232):60217-60218.

________. 2002a. Extension of Comment Period on the Draft Recovery Plan for

Coastal Plants of the Northern San Francisco Peninsula. Federal Register

67(47):10924-10925.

________. 2002b. Second Extension of Comment Period on the Draft Recovery Plan

for Coastal Plants of the Northern San Francisco Peninsula. Federal Register

67(131):45532-45533.
235

U.S. Fish and Wildlife Service and National Oceanic and Atmospheric Administration. 

1994a. Endangered and Threatened Wildlife and Plants: Notice of Interagency

Cooperative Policy on Recovery Plan Participation and Implementation Under

the Endangered Species Act. Federal Register 59:34272. July 1, 1994.

________. 1994b. Threatened and Endangered Wildlife and Plants: Notice of

Interagency Cooperative Policy for Peer Review in Endangered Species Act

Activities. Federal Register 59:34270. July 1, 1994.

Vila, M., and C. M. D’Antonio. 1998. Hybrid vigor for clonal growth in Carpobrotus 

(Aizoaceae) in coastal California. Ecological Applications 8: 1196-1205.

Wayne, L. 1996. An evaluation of rare plant monitoring (Lessingia germanorum and 

Chorizanthe cuspidata) in the Presidio, GGNRA. Report to Presidio Natural

Resources, Golden Gate National Recreation Area, National Park Service, Fort

Mason, San Francisco, California.

Webster, G. L. 1993. Euphorbiaceae (including Croton). Pp. 567-577 in The Jepson

manual: higher plants of California, ed. J. C. Hickman. University of

California Press. 1,400 pp.

Wells, P. V. 1968. New taxa, combinations, and chromosome numbers in

Arctostaphylos (Ericaceae). Madroño 19:193-210.

________. 1991. The naming of the manzanitas. The Four Seasons 8:46-70.

________. 1993. Arctostaphylos, pp. 545-559 in The Jepson manual: higher plants

of California, ed. J. C. Hickman. University of California Press. 1,400 pp.

________. 1998. New combinations in Arctostaphylos (Ericaceae): annotated list of

changes in status. Madroño 35:330-341.

White, P. S. 1996. Spatial and biological scales in reintroduction. 49-86 in Falk, D.,

C. I. Millar, and M. Olwell. 1996. Restoring Diversity: Strategies for

Reintroduction of Endangered Plants. Island Press, Washington, D.C. 
236

Wiedemann, A. M. 1966. Contributions to the plant ecology of the Oregon coastal

sand dunes. Ph.D. dissertation, Oregon State University, Corvallis, Oregon.

255 pp.

Wilkin, D. H. Silene. Pp. 488-493 in The Jepson manual: higher plants of

California, ed. J. C. Hickman. University of California Press. 1,400 pp.

Wood, M. 1996. Rare and endangered plants of San Francisco’s wild and scenic

places, 1st ed. California Native Plant Society, Yerba Buena Chapter.

B. Personal Communications

Albert, Marc. 1996-1998. Golden Gate National Recreation Area, National Park

Service, Natural Resources Branch, Presidio, San Francisco, California.

Brastow, Peter. 1999. Golden Gate National Parks Association, San Francisco,

California. [Please note that the Golden Gate National Parks Association has

changed its name to the Golden Gate National Parks Conservancy.]

Cannon, Joe. 1996-1998. Golden Gate National Recreation Area, National Park

Service, Natural Resources Branch, Presidio, San Francisco, California.

Chasse, Michael. 1998-1999. Golden Gate National Parks Association, San

Francisco, California.

Coulter, Malcolm. 1997. Point Reyes Bird Observatory, Bolinas, California

Elam, Diane. 1999. U.S. Fish and Wildlife Service, Sacramento Fish and Wildlife

Office, Sacramento, California.

Farrell, Sharon. 1998-1999. Golden Gate National Recreation Area, National Park

Service, Natural Resources Branch, Presidio, San Francisco, California.

Feyerabend, Kerry. 1998. Presidio Trust, San Francisco, California.
237

Forbes, Holly. 1998-1999. University of California Botanical Garden, Berkeley,

California.

Hatch, Daphne. 1994-1998. Golden Gate National Recreation Area, National Park

Service, Natural Resources Branch, Presidio, San Francisco, California.

Holloran, Peter. 1998-2001. California Native Plant Society, Yerba Buena Chapter,

San Francisco, California.

Hornor, Nancy. 1996. Golden Gate National Recreation Area, National Park Service,

San Francisco, California.

Mahoney, Donald. 1998-1999. Strybing Arboretum, San Francisco, California.

Markos, Staci. 1998. University of California at Berkeley, Berkeley, California.

Mencke, Mark. 1998. San Francisco State University, San Francisco, California

Moritsch, Barbara. 1999. Plant Ecologist, Point Reyes National Seashore, California.

Petrilli, Mary. 1997-1998. Golden Gate National Recreation Area, National Park

Service, San Francisco, California.

Sigg, Jacob. 1998. California Native Plant Society, San Francisco, California.

Smith, Susan M. 1998. San Francisco, California.

Vasey, Michael. 1998-1999. San Francisco State University, San Francisco,

California.

Wadsworth, Monica. 1997-1999. Golden Gate National Parks Association, Fort

Mason, San Francisco, California.

Wood, Michael. 1999. California Native Plant Society, Yerba Buena Chapter, San

Francisco, California.
238

C. Unpublished References

Baye, Peter. 1990 to 1999. Unpublished field notes. San Francisco, California.

Connors, Peter. 1998. Vascular plant species list for Bodega Head and Bodega dunes. 

Bodega Marine Laboratories, University of California, Davis. 6 pp. 

Gaar, Greg. 1999. Private collection of historic San Francisco photography, San

Francisco.

Golden Gate National Recreation Area, National Park Service, Fort Mason, San

Francisco), unpublished file information.

U.S. Army Corps of Engineers, San Francisco District. Unpublished file information

and historic aerial photographs of San Francisco.

Whelchel Adam. 1998. Memorandum: insect visitor list for Lessingia germanorum.

San Francisco State University. 2 pp.

U.S. Fish and Wildlife Service. Endangered species file information on Raven’s

manzanita, Arctostaphylos hookeri ssp. ravenii, sheets dated 1987.
239

D. In Litt. References

Markos, Staci. 2002. Peer review letter on draft recovery plan dated April 23, 2002. 

Jepson Herbarium and Department of Integrative Biology, University of

California; Berkeley. 2pp.

Middleton, Craig. 2002. Comment letter on draft recovery plan dated May 14, 2002. 

Presidio Trust; San Francisco, California. 20pp.

O’Neill, Brian. 2002. Comment letter on draft recovery plan dated September 9,

2002. National Park Service, Golden Gate National Recreation Area; San

Francisco, California. 24pp.
Appendix I - 1 

VIII. APPENDICES

APPENDIX I. Native Vascular Plant Species of Coastal Dunes and Dune Slacks

of the Northern San Francisco Peninsula

The following list of vascular plants recorded from the San Francisco dune system is

compiled from Howell et al. (1958) Brandegee (1892), Thomas (1961), and recent

field observations (P. Baye unpubl. data 1984 to 1999). Locations are cited from

Howell et al. (1958) unless otherwise noted. Species included are either currently

verified from dune habitats, or have been identified on “sandy soils” (older organicstained, stabilized dune scrub or grassland) or “sandy flats” (dry dune slack, deflation

plain) at locations known to include recent (Holocene) dune deposits, and are

presumed to be dune localities. Some sandy soil locations near Lake Merced or Ocean

View may be Colma formation sands rather than dune, however. Wetland plants

collected from localities of historic dune slacks are included. 

Nomenclature generally follows Hickman (1993), based on synonymy given in that

volume or Munz (1959), unless infraspecific taxa are cited that are not recognized in

Hickman (1993). Question marks (?) indicate some uncertainty of collection on dune

substrates, or somewhat uncertain taxonomic status. Localities cited in Howell et al.

(1958) named with “obligate” (high fidelity or frequency) dune species or multiple

collections of dune species include Point Lobos, Lobos Creek, Presidio, Lone

Mountain, “dunes, Sunset district,” Sunset Heights, Lake Merced, Golden Gate Park,

Ocean beach, Baker Beach/above Baker Beach, and near Cliff House. Conventional

common names are compiled primarily from Best et al. 1996 and Hickman 1993. 

Arbitrary English names are given in brackets [ ] when no conventional English name

is known.

PTERIDOPHYTES

(Ferns and fern-like plants)

AZOLLACEAE

Azolla filiculoides Lam. AMERICAN WATER-FERN, MOSQUITO-FERN. “...among

dunes in Sunset district...”. Localities undoubtedly refer to perennially ponded dune slacks. 
Appendix I - 2 

EQUISETACEAE

Equisetum hyemale ssp. affine (E. hyemale “var. elatum (Engelm.) Morton”) GIANT

SCOURING RUSH. “moist or wet places on coastal dunes...Lobos creek..Lake Merced”. 

Also occurs on sand slope with dune vegetation at Hillside Park, Daly City.

POLYPODIACEAE

Polypodium californicum Kaulfuss. CALIFORNIA POLYPODY FERN, CALIFORNIA

SHIELD FERN. “...rarely sandy slopes of hills...above Bakers beach...Land’s End...Lone

Mountain....dunes south of Golden Gate Park...”. Also rare on old stable dunes of a northfacing slope at Fort Funston.

Pteridium aquilinum (L.) Kuhn var. pubescens L. Underw. BRACKEN FERN. 

“widespread and rather common on dune hills...Presidio, Lone Mountain, Golden Gate Park,

Sunset Heights”. Common in stable dune grassland and scrub in San Francisco today.

ANGIOSPERMS (Anthophytes)

Flowering Plants

Dicots (Broadleaf flowering plants)

APIACEAE

Daucus pusillus Michaux RATTLESNAKE WEED. “brushy slopes and in open

grassland...Sunset Heights...Lake Merced”. Locally common in stable dunes of the Presidio,

Sunset Heights, and Fort Funston.

? Oenanthe sarmentosa Presl. WATER CELERY. This species likely occurred in some wet

dune slacks of the Sunset and Richmond districts; it was recorded in streams and lakes

adjacent to or within dune systems, such as Lobos Creek and Lake Merced.

ASTERACEAE

Achillea millefolium L. (A. borealis Bongard ssp. arenicola (Heller) Keck) YARROW. 

“..common in...sandy soil in natural areas...sand hills, San Francisco...Presidio...Lone

Mt...dunes, Sunset district...Lake Merced....”. Still locally common.

Agoseris apargioides (Less.) E. Greene var. eastwoodiae COAST DANDELION. “Common

on dunes, grassy hills...sand hills near San Francisco...Presidio...Point Lobos...Lone

Mountain....dunes, Sunset District....”. Still locally common.

Ambrosia chamissonis (Less.) E. Greene. (Franseria chamissonis Lessing; varieties

bipinnatisecta and chamissonis not recognized; both forms and intermediates occur in San

Francisco) BEACH-BUR. “Sandy flats, occasionally in clayey soil near the shore: Presidio;

Lone Mountain...Golden Gate Park; dunes, Sunset district...Lake Merced...Fort Mason....near

Islais Creek.”. A dominant species of foredunes, mobile dunes, and early phases of stable

dunes. Also frequent at Ocean Beach.
Appendix I - 3 

Anaphalis margaritacea (L. ) Benth. & Hook. PEARLY EVERLASTING. “brushy

slopes....dunes, Sunset district...Lone Mountain....Presidio...”

Artemisia pycnocephala De Candolle DUNE SAGEWORT, DUNE SAGE. “Dunes and

sandy flats: Lobos Creek...Bakers Beach..Laurel Hill Cemetery...Lake Merced.” Also occurs

at Sunset Heights. San Francisco is the type locality. 

Baccharis pilularis DC. COYOTE BRUSH. “Dune flats and open hillsides...Presidio...Point

Lobos...Lone Mountain....dunes, Sunset district.... Lake Merced...”. Still a dominant species

of dune scrub locally.

Cirsium occidentale (Nutt.) Jepson var. occidentale. COBWEBBY THISTLE, WESTERN

THISTLE. “Common on sandy slopes or dune flats, rarely in rocky soil...above Bakers

Beach...Golden Gate Park...Sunset Heights....Ocean View....Lake Merced...”. Persists at

Baker Beach, Fort Funston dunes. Funston plants are notably short and basally branched,

possibly tending towards var. compactum, but are distinctly cauline.

Cirsium occidentale (Nutt.) Jepson var. compactum Hoover COMPACT COBWEBBY

THISTLE. Taxonomic affinity of San Francisco specimens is questionable. Apparently

extirpated in San Francisco (Skinner & Pavlick 1994, Wood 1996), probably always rare here

if referable to this taxon. Formerly not distinguished from var. occidentale. 

Ericameria ericoides (Less.) Jepson (Haplopappus ericoides (Less.) Hook.& Arn.) FALSE

HEATHER. “common in low shrub of dunes and sandy hills: Presidio...Lands End...Lone

Mountain....Golden Gate Park....dunes, Sunset district...Lake Merced....Chamisso collected the

type in San Francisco....”. Persists at Sunset Heights, Baker Beach, Presidio, Fort Funston

(“Lake Merced”) localities.

Erigeron glaucus Ker-Gawler SEASIDE DAISY. “Coastal bluffs, sandy flats, and rocky

hills: Presidio....west of Lake Merced [Fort Funston]...”. Found locally primarily on older

stable dunes.

Erigeron foliosus Nutt. var. franciscensis G. Nesom. LEAFY DAISY. “Grassy slopes near

brush...not common; Point Lobos...Lake Merced...”. Recorded generally from habitats

including dune grassland (Hickman 1993). Not reported recently from local dunes; possibly

extirpated.

Eriophyllum staechadifolium Lascaga (E. s. var. artemisiaefolium (Lessing) Macbride) 

LIZARD-TAIL, SEASIDE WOOLY-SUNFLOWER. “Bluffs and brushy hills on or near the

coast: ....Golden Gate Park...Lake Merced...”. Occasional on older dunes today at Fort

Funston, Baker Beach.

Gnaphalium stramineum Kunth (Gnaphalium chilense Sprengel) CUDWEED. “Common

and widespread in sandy and clayey soils in grassland or brush: Presidio...Golden Gate

Park...dunes, Sunset district...”. Occasional to locally common in San Francisco dunes.

Gnaphalium purpureum L. CUDWEED. “Rather common in open grassy places, rarely

ruderal: Presidio...Lone Mountain...Golden Gate Park...Lake Merced...”. Occasional to

locally common in older San Francisco dunes.
Appendix I - 4 

Grindelia hirsutula Hook. & Arn. var. maritima (E. Greene) M.A. Lane (G. maritima

(Greene) Steyermark) SAN FRANCISCO GUMPLANT. “Coastal bluffs, sandy flats, and

open or brushy slopes...Presidio....Lake Merced...probably the San Francisco Grindelia

collected by Chamisso in 1816....” Occurs at Fort Funston on wind-reworked Merced

formation sands, possibly planted.

Grindelia stricta DC. var. platyphylla (E. Greene) M. A. Lane (G. arenicola Steyermark var.

pachyphylla Steyermark) [BROADLEAF GUMPLANT] . “A collection made by L.S. Rose

in 1930 is cited from San Francisco by Steyermark (Ann. Mo. Bot. Gard. 21:596, -1934).” 

This species, regionally found in dunes at Point Reyes, was not cited for the San Francisco

flora by Brandegee (1892), and probably was rare here. Common on bluffs of San Mateo

coast and Point Reyes dunes and bluffs. Now extirpated in San Francisco.

Lasthenia glabrata Lindley ssp. glabrata SMOOTH GOLDFIELDS. “Wet valley lands

....reported from the western part of the city across to Visitacion Valley”. “Wet valleys” in the

western part of the city are likely to have included dune slacks since the western part of the

city was almost entirely covered by the San Francisco dune sheet.

Lasthenia minor (Baeria minor (DC.) Ferris) GOLDFIELDS. “Moist places in sandy and

clayey soil...Lake Merced...Presidio golf course...”. Though not specifically cited from San

Francisco dunes, “moist places in sandy...soil” at Lake Merced and the Presidio golf course”

are likely former dune occurrences. This species is locally common on older dunes of Point

Reyes and Dillon Beach, Marin county, and would have been a likely species of dune slacks

and old dunes in San Francisco.

Layia carnosa (Nutt.) Torrey & A. Gray BEACH LAYIA. Thomas (1961) cites this species

as “occ. on coastal sand dunes, known so far locally only from San Francisco”. No collections

are cited. Not reported in Howell et al. 1958, or in California collections of California

Academy of Sciences, University of California, or Jepson Herbarium. Probably occurred at

most as a local small population since it was not recorded by Brandegee (1892) or Howell et

al. 1958. Now extirpated; nearest population is at Point Reyes, in dunes near Abbotts Lagoon.

Layia platyglossa (Fischer & C. Meyer) A. Gray TIDY TIPS. “Common in grassy

places...San Francisco sand hills...Presidio....Point Lobos....Sunset Heights...Lake Merced”.

Lessingia germanorum Cham. SAN FRANCISCO LESSINGIA. “Sandy flats and dune

hills: San Francisco...Presidio...Lone Mountain...Lake Merced....The original collection, on

which the genus and species were based, was made by Chamisso in San Francisco in 1816...”. 

(See text and Table 1 of this recovery plan for detailed historic locality information).

Lessingia filaginifolia (Hook. & Arn.) M.A. Lane var. californica (Corethrogyne californica

De Candolle) CALIFORNIA-ASTER. “Reported by Greene...[1894]...from sandy hills, Lake

Merced.” Presumed extirpated here on dunes.

Microseris bigelovii (A. Gray) Schultz-Bip. MICROSERIS. “Widespread and rather common

on sandy, rocky, or clayey soils of open flats and slopes: ...Presidio...Lone Mountain.. ”.
Appendix I - 5 

Senecio aronicoides DC. RAGWORT. “Grassy or brushy slopes, generally in partial shade: 

above Bakers Beach, Presidio...Sunset Heights...Lake Merced...”. Current status on local

dunes uncertain; not recently reported.

Solidago spathulata DC. GOLDENROD. “Open slopes and flats in rocky, clayey, or sandy

soil: ....Lone Mountain...Sunset Heights....Lake Merced....collected....by Chamisso...”. 

Remnant natural dune populations today mostly in Sunset Heights; scarce in the Presidio

dunes.

Tanacetum camphoratum Less. DUNE TANSY. “ Forming colonies on sandy flats and

dunes: San Francisco sand hills...Point Lobos....dunes, Sunset District...Ocean beach...type

collected in San Francisco by Chamisso.”. Remnant natural dune populations in the Presidio,

dunes near Cliff House (above Balboa at Great Highway), Sunset Heights, Fort Funston. 

Nearest population is Dillon Beach, Marin County.

Uropappus lindleyi (DC.) Nutt. (Microseris linearifolia (DC.) Schultz-Bip.) SILVERPUFFS. “Grassy or brushy slopes....Lobos Creek, Presidio...dunes, Sunset district....Lake

Merced...”. Common in grassland and ruderal habitats as well as on dunes.

BORAGINACEAE

Amsinckia spectabilis Fischer & C. Meyer COAST FIDDLENECK. “common on dunes and

on sandy and grassy flats: above Bakers Beach...Point Lobos..Lone Mountain...dunes, Sunset

district...Lake Merced”. Persists in Presidio dunes.

(?) Cryptantha clevelandii E. Greene (C. hispidissima E. Greene, C. c. var. florosa. I.M.

Johnston) [CLEVELAND’S POPCORNFLOWER]. “sandy slopes and flats...San

Francisco....Lake Merced”. May be locally extirpated.

Cryptantha leiocarpa (Fischer & Meyer) E. Greene DUNE POPCORNFLOWER. “on dunes

and sandy flats....near Bakers Beach....Presidio...Golden Gate Park...”. Persists at historic

localities and restored dunes nearby.

? Nemophila pedunculata Douglas ex Benth. [STALKED NEMOPHILA]. “...open grassland

or under brush....Point Lobos...Lone Mountain...Lake Merced”. Probably extirpated in San

Francisco dunes. Occurs in stable dunes in Monterey Bay, Morro Bay.

? Plagiobothrys spp. (P. chorisianus (Allocarya chorisiana), P. reticulatus var. rossorianum

(Allocarya diffusa), possibly other spp., like P. stipitata, may have occurred locally in dune

slacks. POPCORNFLOWERS. They are cited from historic localities that included dunes and

possibly dune slacks (Lake Merced, near Lobos Creek, Golden Gate Park. [uncertain. cited as

“locally common in moist places,” including Lobos Creek and Golden Gate Park. Possibly

present in dune slacks; Plagiobothrys spp. (within former genus Allocarya) are rarely found at

Point Reyes dune slacks, and are locally abundant at wet young slacks at Manchester Beach

dunes near stream mouths. 
Appendix I - 6 

BRASSICACEAE

Erysimum franciscanum Rossbach FRANCISCAN WALLFLOWER. “...dune hills and

ocean bluffs in the western part of the city...sand hills near San Francisco...above Baker

Beach...Lone Mountain...Sunset Heights...”. Persists at Baker Beach, Presidio, Fort Funston

and Sunset Heights dunes.

CARYOPHYLLACEAE

Minuartina californica (A. Gray) Mattf. (Arenaria californica (A. Gray) Brewer) 

CALIFORNIA SANDWORT. “sandy places and grassland.... ‘meadows among sand dunes

south of the Park’, acc. Brandegee.”.

Arenaria paludicola Robinson MARSH SANDWORT. “Swamps....Fort Point.” Long

extinct; last reported by Hans Behr in the 1850's from the Presidio Swamp, located behind or

within a barrier beach and dune complex, now Crissy Field.

Cardionema ramossisimum (J.A. Weinm.) Nelson & J. MacBr.) SAND MAT. “dune

hills...Bakers Beach...dunes near Cliff House.. dunes, Sunset district....”. Persists at these

localities; also at Sunset Heights, Fort Funston.

Cerastium arvense L. FIELD CHICKWEED. “Sunset Heights...Lake Merced”. Persists at

Sunset Heights dune remnants.

(?) Sagina maxima A. Gray var. crassicaulis (S. Watson) G. Crow (S. crassicaulis) BIG

PEARLWORT. “moist bluffs and flats just above the ocean: Bakers Beach...Point Lobos”. 

Presumed extirpated in dunes and bluffs in San Francisco. This species is primarily found at

sandstone bluffs, but occurs in dune slacks at Point. Reyes; “flats just above the ocean” at

Baker Beach probably refers to dune slacks. S. occidentalis also possibly occurred in dune

slacks.

Silene verecunda S. Watson ssp. verecunda SAN FRANCISCO CAMPION. “above Bakers

Beach...brushy dunes, Sunset district”. San Francisco is the type locality. Persists at Baker

Beach; reintroduced to Lobos dunes, Presidio.

Stellaria littoralis Torrey COAST STARWORT. “In wet sandy soil of maritime slopes: 

Lands end...”. Extirpated in San Francisco. Locally abundant in dune slacks of Point Reyes

and Brazil Beach (Tomales Bay mouth).

CHENOPODIACEAE

Chenopodium californicum (S. Watson) S. Watson. CALIFORNIA GOOSEFOOT. “Rare

on brushy hillsides: coastal slopes, Presidio...sandy hollow east of Lake Merced...”. 

Occasional in old, weathered stable dune soils, dune scrub of Monterey Bay, Morro Bay, and

Point Reyes; collected by M. Chassez, Golden Gate National Parks Association in 1997, near

Rob Hill, Presidio, on old stable dune soil.
Appendix I - 7 

Atriplex californica Moquin CALIFORNIA SALTBUSH. “...maritime species of bluffs and

dune flats: Point Lobos...Fort Point according to Brandegee”. Extirpated in “dune flats”;

persists in bluffs at Point Lobos and west of Fort Point. Uncommon to rare on San Mateo

coast bluffs; the nearest populations are in Pacifica.

Atriplex leucophylla (Moquin) D. Diedrich. BEACH SALTBUSH. “Sandy shores along bay

and ocean”. Still frequent some years at Ocean Beach, occasionally Crissy Field.

CONVOLVULACEAE

Calystegia soldanella (L.) R. Br. BEACH MORNING-GLORY “..sands near the shore of the

Presidio” according to Brandegee. Extirpated in San Francisco, but reintroduced to Crissy

Field (Presidio) in 1998 from Linda Mar (Pacifica) and Half Moon Bay.

CRASSULACEAE

Crassula connata (Ruiz Lopez & Pavon) A. Berger. (Tillaea erecta Hook. & Arn.) SAND

PYGMY WEED. “sandy flats...near Baker Beach...Land’s End...Lone Mountain...”. Persists

in dunes near Baker Beach.

Dudleya farinosa Abrams (Escheveria farinosa Lindley) COAST DUDLEYA. Occurs as

locally frequent component of vegetation on old stable remnant dunes of Sunset Heights and

Fort Funston. No dune collections are cited in Howell et al. 1958, who described only bluff

and rocky outcrop habitats. 

CUCURBITACEAE

Marah fabaceus (Naudin) E. Greene CALIFORNIA MAN-ROOT. “Common on

dunes.....Lands End to Fort Point...near Lobos Creek...above Bakers Beach....dunes, Sunset

district...” Persists in dunes near Baker Beach, Sunset Heights.

EUPHORBIACEAE

Croton californicus Muell. CALIFORNIA CROTON. “open sandy flats or sandy soil on

brushy slopes...Presidio....Lobos Creek....above Bakers beach...Lone Mountain...Lake

Merced.”. Extirpated near Lake Merced (Fort Funston dunes) and Lone Mountain, but persists

at Sunset Heights (Hawk Hill) and Presidio dune localities. San Francisco is the type locality

and northern coastal range limit.

FABACEAE

Astragalus nuttallii (Torrey & A. Gray) J. Howell var. virgatus (A. Gray) Barneby. 

[NUTTALL’S MILKVETCH]. “Brushy places, usually on sandy soil: Presidio...Point

Lobos...Ocean View...Lake Merced...”. Likely occurred on sandy bluff (Merced, Colma

sandstone) or Holocene dune soils. Occurs locally in restored dunes at Fort Funston (Lake

Merced).

Lathyrus littoralis (Nutt.) Endl. BEACH PEA. “Dunes and ocean beach west of Lake

Merced [Howell, Raven]”. Brandegee (1892) indicated the same population south of the
Appendix I - 8 

outlet of Lake Merced. California Native Plant Society, Yerba Buena Chapter, located a new

population on graded sands within the water treatment facility at this locality, in 1998 (P.

Holloran pers. comm. 1998).

(?) Lathyrus vestitus Nuttall var. vestitus (L. v. ssp. bolanderi Watson. HILLSIDE PEA. 

“...grassy and brush slopes: Near Baker Beach..Sunset Heights...Lake Merced...”. Not

currently reported from dune sand substrates.

Lotus heermanii (Durand & Hilgard) E. Greene var. orbicularis (A. Gray) Islay ( L.

eriophorus Greene, Hosackia tomentosa Brewer & Watson) WOOLLY LOTUS. “Sandy flats

and slopes: dunes, Sunset district...Lake Merced...”. Persists at least at one locality at Fort

Funston.

Lotus humistratus E. Greene SHORT-PODDED LOTUS. “ ...‘common in the western part’

[of the city]...” according to Brandegee (1892).

Lotus scoparius (Nuttall) Ottley DEERWEED, CALIFORNIA BROOM. “Sandy

flats...Presidio...Point Lobos...Sunset Heights...dunes, Sunset district...Lake Merced..”. Still

abundant locally in dune scrub.

Lotus strigosus (Nuttall) E. Greene BISHOP’S LOTUS. “Sandy flats and bluffs: Lobos

Creek...Lone Mountain. ...dunes, Sunset district...”.

Lotus wrangelianus Fischer & C. Meyer ( L. subpinnatus Lagasca misappl.) CHILE LOTUS.

“sandy ...soil ....Presidio...Lone Mountain.”.

Lupinus arboreus Sims. YELLOW LUPINE, BUSH LUPINE, TREE LUPINE. “Common

and widespread in sandy or clayey soil, frequently growing in disturbed places: dunes, Sunset

District Rubzoff...”. Not recorded on dunes in earliest local surveys of woody vegetation by

Bolander (1863). Actively planted for dune stabilization (McLaren 1924, Clary 1980). By

late 1900's, Brandegee (1892) recorded L. arboreus as frequent in sands of western half of

city. San Francisco is the probable source of seed that grew the type specimens. No blue

(“var. eximius”) types occur in San Francisco.

Lupinus bicolor Lindley MINIATURE LUPINE. “Widespread in sandy..soil...our

commonest annual lupine...above Bakers Beach...Lone Mountain...dunes, Sunset district....”. 

Now scarce in San Francisco; persists at Sunset Heights dune remnants and around Lobos

Dunes.

Lupinus chamissonis Eschscholtz BLUE BEACH LUPINE, SILVER BEACH LUPINE,

CHAMISSO’S LUPINE. “Sandy soil of dune hills and flats....: above Bakers

Beach...Presidio...dunes, Sunset district..Lake Merced...”. Type locality is San Francisco, in or

near Presidio. Still locally abundant at most remnant dune sites other than Crissy Field.

Lupinus nanus Douglas ex Bentham. SKY LUPINE. “Hillsides and flats in sandy and clayey

soil: sand hills near San Francisco...Presidio..Lone Mountain...Lake Merced...”. Now scarce

in dunes; spontaneous occurrences are doubtful.
Appendix I - 9 

Lupinus variicolor Steudel. [COAST LUPINE]. “Open grassland in sandy or clayey soil....”. 

Occurs primarily on coastal bluffs. Current spontaneous dune localities uncertain.

(?) Trifolium macraei Hook. & Arn. CHILE CLOVER. “Sandy ....open hillsides and flats,

rather common...Presidio...” Probably occurred in dune slacks, as at Dillon Beach, Marin

County.

(?) Trifolium wildenovii Sprengel (T. tridentatum Lindley) TOMCAT CLOVER. 

“Widespread in open grassy places... Lone Mountain....Presidio...Sunset Heights...Lake

Merced...”.

(?) Trifolium wormskioldii Lehmann COW CLOVER. “wet meadows and hillside

seepages... Presidio...Pt. Lobos...Lone Mountain....”. Probably occurred in some moist dune

slacks, as at Dillon Beach, Marin County.

FAGACEAE

Quercus agrifolia Nee COAST LIVE OAK. “...dune hills...Lobos Creek...Golden Gate

Park...”. Persists in old remnant dunes behind Marine Hospital and Lobos Creek banks,

Presidio, and near the Fuchsia Dell, Golden Gate Park.

GENTIANACEAE

(?) Centaurium spp. CENTAURY. Centaurium davyi (Jepson) Abrams (possibly C.

trichanthum Grisb.) Robinson) may be expected in dune slacks and old dune soils, as at Pt.

Reyes, Marin County, and Tenmile dunes, Mendocino County. Howell et al. (1958) state that

the identity of “C. douglasii” of Behr’s San Francisco flora is uncertain. C. davyi occurs

locally on coastal headland grassland with thin old dune soils at Pacifica. C. muehlenbergii

occurs on clayey soils in southern San Francisco, but is not generally found on dunes.

(?) Cicendia quadrangularis (Lam.) Grisb. (Microcala q. (Lam.) Grisb.) AMERICAN

MICROCALA. This species was reported by Brandegee from Presidio. It occurs in the Bay

area in seasonal wetlands and vernal pools. It possibly occurred here in sparsely vegetated

dune slacks, as at Tenmile Dunes, Mendocino County.

HYDROPHYLLACEAE

Phacelia californica Cham. CALIFORNIA PHACELIA. “...common....sandy soil..Sunset

Heights...dunes, Sunset district...”. Persists in relict old dunes at Sunset Heights. Otherwise

found primarily on stony soils and bluffs. Occasional in old stable dunes of the central coast.

Phacelia ciliata Benth. GREAT VALLEY PHACELIA. “sandy flats...near Lone

Mountain...”. Probably an occasional or incidental species in local dunes. Current status in

San Francisco dunes uncertain.

Phacelia distans Benth. COMMON PHACELIA, WILD HELIOTROPE. “common on sandy

flats....Presidio....Point Lobos....Lone Mountain...dunes, Sunset district....”. Still locally

abundant in stable dunes and other plant communities.
Appendix I - 10 

LAMIACEAE

Monardella undulata Benth. WAVY-LEAVED MONARDELLA. “sandy flats at Lake

Merced”. It is not certain whether this locality was east (on Colma formation sand deposits) or

west (on recent surface dunes) of Lake Merced. The species occurs primarily on dune sand in

most of its range. The nearest population is at Point Reyes dunes.

Monardella villosa Benth. ssp. franciscana (Elmer) Jokerst FRANCISCAN COYOTEMINT. “sandy slopes...more common in western part of the city: Lake Merced...”. Not

recently reported; may be extirpated locally in dunes.

Satureja douglasii (Benth.) Briq. YERBA BUENA. “brushy slopes...dunes, Sunset district”. 

Occurs marginally in moist, shaded dune slopes.

MYRICACEAE

Myrica californica Cham. CALIFORNIA WAX-MYRTLE. “in wet and marshy places in

gullies and on dunes....dunes, Sunset District”. Occurs also in dune slacks of Marin, Sonoma,

and Mendocino Counties. San Francisco is the type locality.

ONAGRACEAE

Clarkia davyi (Jepson) H. Lewis & M. Lewis DAVY’S CLARKIA. “Sandy flats along the

coast: dunes, Sunset district...Lake Merced ...southernmost station for the species.”

(?) Clarkia rubicunda Lindley (H. Lewis & M. Lewis) CLARKIA. “Locally common on

grassy or brushy slopes and coastal bluffs...hills near Golden Gate Park.” This species occurs

on Pleistocene sand of Daly City, at San Bruno Mountain (McClintock et al. 1990), and Fort

Funston.

Camissonia cheiranthifolia (Sprengel) Raim ssp. cheiranthifolia (Oenothera cheiranthifolia

Hornemann) BEACH EVENING-PRIMROSE. “Common on sandy slopes and flats: above

Bakers Beach...Richmond district...Lone Mountain...Golden Gate Park...dunes, Sunset

district...Lake Merced...”. Common in nearly all dune remnants with vegetation gaps. The

subspecies C.c. ssp. suffruticosa was introduced to Crissy Field dunes in a seed mix from

obtained from southern coastal California (Terri Thomas pers. comm. 1997), and has spread

there and to Fort Funston (through transplantation), where it has apparently hybridized with

native populations to form intermediates (petal sizes exceeding the range of native subspecies,

coarser habit).

Camissonia contorta (Douglas) Raven (Oenothera contorta Douglas ex Hooker var.

strigulosa (Fischer & Meyer) Munz, misapplied). [ TWISTED EVENING-PRIMROSE]. Not

distinguished from C. strigulosa (as Oe. contorta var. strigulosa; see below) in Howell et al.

(1958). Status in San Francisco is unclear; typically found inland.

Camissonia micrantha (Sprengel) Raven (Oenothera micrantha Hornemann) [SMALLFLOWERED EVENING-PRIMROSE] “Coastal bluffs, dunes, and sandy flats: above Bakers

Beach....near Point Lobos..Golden Gate Park...Lake Merced...”.
Appendix I - 11 

Camissonia strigulosa (Fischer & Meyer) Raven (Oenothera contorta Douglas ex Hooker var.

strigulosa (Fischer & Meyer) Munz). [TWISTED EVENING-PRIMROSE] “Dunes and

sandy flats: above Bakers Beach...near Mountain Lake...Richmond district...Lone

Mountain....Golden Gate Park...dunes, Sunset District...Lake Merced.”

Oenothera elata Kunth ssp. hookeri (Torrey & A. Gray) W. Dietr & W.L. Wagner (Oe.

hookeri Torrey & A. Gray) “...brushy slopes and moist places about Lake Merced...”. Occurs

today in Fort Funston dunes, particularly northeast-facing slopes with dune scrub.

NYCTAGINACEAE

Abronia latifolia Eschscholz YELLOW SAND VERBENA. “dune hills and sandy

flats...Bakers Beach...Lands End...near Cliff House...dunes, Sunset...coastal dunes south of

Fort Funston...”. The species is based on a San Francisco collection made on the second

Kotzebue expedition in 1824. A dominant species of foredunes and early phases of stable

dunes. Occurs also at Ocean Beach foredunes.

Abronia umbellata Lamarck ssp. umbellata (introgressive with A. latifolia) PINK SAND

VERBENA. “Rare on sandy beaches: Crissy Field, Presidio...Hunters Point according to 

Brandegee”. Local light pink to salmon flowered types at Crissy Field are perennial

introgressants, not typical species. Collected by Rubzoff at Ocean Beach (CAS).

PAPAVERACEAE

Eschscholzia californica Cham. CALIFORNIA POPPY. “...dune flats, grassy

hills...widespread and variable: Presidio....Lone Mountain...dunes, Sunset District...”. Persists

at most dune remnants, including Sunset Heights.

Meconella linearis (Bentham) Nelson & MacBride. [ MECONELLA]. “sandy soil of coastal

hills....Lobos Creek....Mountain Lake...Lake Merced”. Not recently reported; may be

extirpated.

Platystemon californicus Benth. CREAM CUPS. “...dunes...San Francisco..”. Not recently

reported from dune remnants.

PLANTAGINACEAE

Plantago erecta E. Morris CALIFORNIA PLANTAIN. “abundant on open grassy slopes and

dune flats, often in shallow soil...above Bakers Beach..dunes, Richmond district...dunes,

Sunset District...Lone Mt.”. Persists at Presidio dunes.

PLUMBAGINACEAE

Armeria maritima (Miller) Willd. var. californica (Boiss.) Pors. SEA-PINK, THRIFT. 

“bluffs, dunes....Point Lobos...Richmond district...Sunset Heights...dunes, Sunset district.” 

Persists in local abundance at Sunset Heights.
Appendix I - 12 

POLEMONIACEAE

Gilia capitata Sims ssp. chamissonis (E. Greene) V. Grant DUNE GILIA. “common and

colonial on dunes and sandy flats...near Bakers Beach...Presidio...Lone Mountain...dunes,

Sunset district...”. Persists at most remnant dunes.

Linanthus grandiflorus (Benth.) E. Greene LARGE-FLOWERED LINANTHUS. “open

grassland; near Presidio ....(near Lake Merced)...”. Occurs locally in old stable dunes at Point

Reyes.

Linanthus parviflorus (Benth.). E. Greene ( L. androsaceus (Benth.) E. Greene ssp. croceus

(Milliken) Mason ) COMMON LINANTHUS. “forming colonies in sandy

soil...Presidio..Lake Merced...”. Not recently reported from local dunes.

Navarretia squarrosa (Esch.) Hook. & Arn. SKUNKWEED. “common on sandy and clayey

flats....Lobos Creek to Fort Point..Presidio...dunes, Sunset District; Lake Merced....”. The

Presidio was the type locality. Scarce in dune remnants; persists very locally behind the

Marine Hospital, Presidio.

POLYGONACEAE

Chorizanthe cuspidata S. Watson. (includes var. marginata Goodman, not recognized) SAN

FRANCISCO SPINEFLOWER. “sandy flats, slopes, and dunes...Bakers Beach, near Cliff

House, Richmond...Lake Merced....”. San Francisco is the type locality. Persists at most dune

remnants with vegetation gaps.

Eriogonum latifolium Smith COAST BUCKWHEAT. “sandy soil...common,

variable...ocean beach....dunes, Sunset”. Persists at most dune remnants, locally abundant.

Polygonum paronychia Cham. & Schldl. DUNE KNOTWEED. “sandy flats and

slopes...above Bakers Beach...Lone Mountain.. Sunset Heights...Ocean beach”. San Francisco

is the type locality. Persists widely, but sparsely, in remnant dunes. Also occurs near Cliff

house, Fort Funston.

Pterostegia drymarioides Fischer & Meyer [PTEROSTEGIA] “...rarely in the open on clayey

or sandy soil...”. Occurs at Baker Beach dunes; probable at Fort Funston.

Rumex maritimus L. (Rumex fueginus Phillipi) GOLDEN DOCK. “...dunes, Sunset

district...”. Likely in dune slacks with bare sand, as in Marin County.

Rumex salicifolius J.A. Weinm. var. crassus (Rech.f.) J. Howell. WILLOW DOCK. “sandy

soil on slopes and flats near the ocean...Bakers Beach...Point Lobos...Ocean Beach west of

Lake Merced”.

PORTULACACEAE

Calandrinia ciliata (Ruiz Lopez & Pavon) de Candolle RED MAIDS. “occasional in

open...sandy...”. Not recently reported in local dunes. Occurs in Monterey Bay dunes,

including those recently stabilized. 
Appendix I - 13 

Claytonia perfoliata Willd. ssp. perfoliata (Montia perfoliata (Donn.) Howell) MINER’S

LETTUCE. “dunes, Sunset district...” Also abundant at Lobos Creek and Bakers Beach

dunes, particularly on moist, shaded, or northern slopes.

RANUNCULACEAE

Ranunculus californicus Benth. CALIFORNIA BUTTERCUP. “....dunes..widespread and

variable....Presidio”.

ROSACEAE 

Acaena pinnatifida Ruiz Lopez & Pavon var. californica (Bitter) Jepson (A. californica

Bitter) ACAENA. “Sandy....flats....Presidio; Point Lobos; Lone Mt....Sunset Heights”.

Fragaria chiloensis (L.) Duchesne BEACH STRAWBERRY. “...dunes...above Bakers

Beach...Lands End..near Cliff House...Lone Mt. Sunset Heights...dunes, Sunset district...Lake

Merced...”.

? Horkelia californica Cham. & Schindl. ssp. californica CALIFORNIA HORKELIA. 

“Sunset Heights” [possible in old dune scrub].

Horkelia cuneata Lindley ssp. cuneata (Potentilla Lindleyi Greene) WEDGE-LEAFED

HORKELIA. “Sandy soil near the ocean...Ocean View and Lake Merced...”. Apparently

extirpated in San Francisco dunes.

Horkelia cuneata Lindley ssp. sericea (A. Gray) Keck KELLOGG’S HORKELIA. “Sand

hills near the ocean: Point Lobos and Sunset Heights (acc. Brandegee)..Lake Merced...Ocean

View...”. Extirpated in San Francisco; nearest dune locality of this rare plant is in Daly City.

? Oemleria cerasiformis (Hook. & Arn.) J.W. Landon (Osmaronia cerasiformis Torrey & A.

Gray ex Hook. & Arn.) OSO BERRY. “above Bakers Beach”. Possibly present in old dune

scrub.

? Potentilla anserina L. ssp. pacifica (Howell) Rousi SILVERWEED. “in marshy places and

about seepages..”. Not specifically cited for dunes or dune slacks, but almost certainly must

have been at least locally common in dune slacks of the Sunset district, with Juncus lesueurii,

as in most dune slacks of the central and north coast.

Rubus ursinus (Cham. & Shlecht.) CALIFORNIA BLACKBERRY. “...above Bakers

Beach...Lobos Creek...Lone Mountain...Golden Gate Park...Lake Merced....San Francisco is

the type locality.”. 

RHAMNACEAE

Rhamnus californica Eschscholtz COFFEE-BERRY. “dunes, Sunset district...Lone

Mountain. ...Presidio....According to Dr. Behr (Zoe 2:3 - 1891) most hill tops in the western

part of the city were formerly crowned by an extensive chaparral of robust plants of this

species...The Presidio is the probable type locality of this species...”. Also occurs at Fort

Funston dunes, Sunset Heights dunes.
Appendix I - 14 

SALICACEAE

Salix exigua Nutt. (S. Hindsiana Benth.) WILLOW “...dunes of the Sunset District acc.

Raven...”.

Salix lasiolepis (S. l. var. bigelovii (Torrey) Bebb indistinct acc. Hickman 1993) ARROYO

WILLOW. “in wet places on strands dunes....dunes, Sunset District....sandhills near

Cliffhouse...Lake Merced” . Willow thickets occur in wet dune slacks (e.g. dunes of

Mendocino county), streamsides, (e.g. Lobos Creek) and hillside seeps with climbing dunes

(e.g. Sunset Heights). They also occur on dunes where mobile dunes override slacks (e.g.

Sunset, Kaufeldt 1954; Pescadero dunes).

SCROPHULARIACEAE

Castilleja wightii Elmer (C. latifolia Hook. & Arn. var. wightii (Elmer) Zeile) WIGHT’S

PAINTBRUSH. “...occasional on brushy slopes: Lake Merced....”. Plants with yellow to

orange bracts and strongly glandular stems below the inflorescence, common on the adjacent

San Mateo coast, that are clearly referable to this species have not been recently verified in

San Francisco dunes. Slightly glandular C. affinis occurs in San Francisco dunes, sometimes

identified as C. wightii, but distinct from C. wightii of the San Mateo coast.

Castilleja affinis Hook & Arn. ssp. affinis (as C. latifolia Hook. & Arn. in Howell et al.

1959) INDIAN PAINTBRUSH. “...slopes in coastal brush and grassland....above Bakers

Beach...Lobos Creek to Fort Point..Point Lobos...dunes, Sunset district..Lake Merced...”. 

Many taxonomically ambiguous plants in San Francisco are atypical for C. affinis ssp. affinis,

and have some foliar characters approaching C. latifolia. This species is the locally common

paintbrush of coastal bluff and dunes scrub south of San Francisco. San Francisco and

Monterey Bay have been variously cited as the type locality of C. latifolia. 

Castilleja exserta (A.A. Heller) Chuang & Heckard ssp. latifolia (S. Watson) Chuang &

Heckard. (Orthocarpus purpurascens Benth. var. latifolius S. Watson) BROADLEAF

PURPLE OWL’S-CLOVER. “...Dunes and coastal grassland, often common locally: Lake

Merced...near Ocean View...”. Presumed extirpated here. Locally abundant in parts of San

Bruno Mountain, San Mateo coast (Rockaway Head, Pacifica, on old dune soil), Point Reyes,

and in older Monterey Bay dunes.

Linaria canadensis (L.) Dum.-Cours. (including L.c. var. texana (Scheele) Pennell) BLUE

TOADFLAX. “...forming colonies locally in sandy places: ...near Bakers Beach...dunes,

Sunset district...Lake Merced...”. Found some years near Baker Beach, Lobos dunes.

Limosella aquatica L. MUDWORT. “borders of drying pools and vernally moist hollows: 

Presidio; Lone Mountain....in reporting the station from the Presidio, K. Brandegee (p. 367)

stated, ‘these pools are being filled with sand in the process of grading the tract and will soon

be destroyed.’”. Brandegee’s comment suggests dune slack populations of Limosella; L.

acaulis is present in dune slacks at Point Reyes.

Mimulus guttatus DC (M.g. var. grandis Greene) MONKEY-FLOWER. “...near Bakers

Beach....Lobos Creek....dunes, Sunset district...”. Almost certainly an element of the dune

slack flora, as at Dillon Beach, Marin County.
Appendix I - 15 

Mimulus aurantiacus Curtis STICKY MONKEY-FLOWER. “common on brushy slopes: 

...above Bakers Beach...Lone Mountain....dunes, Sunset District..Lake Merced...”.

Scrophularia californica Cham. & Schindl. CALIFORNIA FIGWORT, BEE-PLANT. 

“common on brushy slopes...Lobos Creek...above Bakers Beach...dunes, Sunset district...”. 

Also locally abundant on remnant dunes of Sunset Heights, Fort Funston.

Triphysaria pusilla (Benth.) Chuang & Heckard (Orthocarpus pusillus Benth.) SMALL

OWL’S-CLOVER. “Rather common on grassy flats and slopes, often in shallow soil: ...near

Bakers Beach...dunes, Sunset district...”.

URTICACEAE

Urtica dioica L. ssp. holosericea (Nutt.) Thorne (U. holosericea Nutt.) HOARY NETTLE. 

“in sandy soil among oaks...Golden Gate Park”. This species also occurs in local abundance at

the margins of dune slacks with organically rich soils.

VIOLACEAE

? Viola adunca Smith. WESTERN DOG VIOLET. “open grassy hills...Bakers

Beach...Sunset Heights...”.

Monocots (grasslike plants, lily-like plants)

CYPERACEAE

Carex obnupta L. Bailey. SLOUGH SEDGE. “wet places among coastal hills and

dunes...Lobos Creek, Golden Gate Park...Lake Merced”. Also known in wet dune slacks at

Point Reyes, Marin County.

(?) Carex pansa L. Bailey. [SAND SEDGE]. Uncertain status in San Francisco dune flora,

from a single questionable record that may have been confused with a Pacific Grove locality. 

Known from dunes in Monterey and San Luis Obispo Counties.

Eleocharis macrostachya Britton SPIKERUSH. “Golden Gate Park...”. Likely in dune

slacks. Found in dune slacks of Point Reyes, Marin County.

(?) Scirpus cernuus Vahl. [SEEP SEDGE] “wet soil of marshes, strands, and seepages”. 

Likely in seasonally ponded or saturated dune slacks.

(?) Scirpus californicus (C. Meyer) Steudel CALIFORNIA TULE. “Golden Gate Park”

(likely component of original wet dune slacks and “Presidio Swamp”).

Scirpus pungens Vahl. (S. americanus Persoon) COMMON THREESQUARE. “...Golden

Gate Park...Lake Merced”. Probable in dune slacks or seasonal streams in dunes, especially

early colonization of saturated sand. Found abundantly in dune slacks of Dillon Beach, Marin

County.
Appendix I - 16 

JUNCACEAE

Juncus bufonius L. TOAD RUSH. “common and widespread in moist or wet soil”. Likely

in seasonally ponded dune slacks, as in central and north coast dune slacks; several likely dune

localities are cited in Howell et al. 1958. 

Juncus lesueurii Bolander. [SALT MARSH RUSH] “wet places on slopes and flats,

sometimes bordering salt marshes” Presidio...Lone Mountain Golden Gate Park”. This

species, and the questionably distinct J. breweri Engelm. and their intermediate forms (not

recognized by Howell et al. 1958), are commonly found in dunes and dune slacks of Marin

County.

Juncus phaeocephalus Engelm. BROWN-HEADED RUSH. “Wet soil of sandy flats or

marshy places, rather common: ....Lone Mountain....Lake Merced...”. Likely in dune slacks,

as at Point Reyes and Dillon Beach, Marin County.

LILIACEAE

Triteleia laxa Abrams (Brodiaea laxa (Bentham) Watson) ITHURIEL’S SPEAR. “dunes in

Sunset District”.

POACEAE

(?) Agrostis pallens Trin. (A. diegoensis Vasey). THIN BENT, DUNE BENT. Described as

“common on open or brushy slopes” by Howell et al. (1958), but not explicitly cited from

dunes. In Marin and Sonoma Counties, this species is commonly found on stable dunes

(Howell 1949, Best et al. 1996), and its occurrence on San Francisco dunes was likely.

Bromus carinatus Hook. & Arn. var. carinatus CALIFORNIA BROME. “dunes, Sunset

District”.

Bromus carinatus Hook. & Arn. var. maritimus Piper) C. Hitchc. (B. maritimus (Piper)

Hitchcock) [MARITIME BROME]. “Forming clumps among coastal dunes and on sandy

hills...Presidio...Point Lobos...Lone Mountain...Sunset Heights...dunes, Sunset District...Lake

Merced”.

Leymus mollis (Trin.) Pilger ssp. mollis (Elymus mollis Trin.) PACIFIC DUNEGRASS,

SEA-RYE. “Dunes along the ocean shore: near cliff house”. Modern populations occur at

Crissy Field beach (dwarfish strains), Sutro baths slopes, Ocean Beach near Irving St., Ortega

to Riviera Streets, and at the north end of Fort Funston dunes.

Leymus pacificus (Gould) E.R. Dewey. [PACIFIC WILDRYE] “Dunes and sandy flats,

where rather frequent... Bakers Beach... Presidio... Sutro Heights...above point Lobos,

Richmond district....Golden Gate Park...”. Also at Fort Funston, dune slope above Balboa at

Great Highway. Some plants intermediate with L. triticoides in some characters.

Leymus x vancouveriensis (Vasey) [VANCOUVER WILDRYE, HYBRID DUNEGRASS] 

“sandy hollow east of Lake Merced”. Extirpated; proposed for reintroduction at Crissy Field

dunes, 1999.
Appendix I - 17 

Festuca rubra L. RED FESCUE. “forming patches locally on ...dunes and coastal

bluffs....above Bakers Beach... Point Lobos... Lone Mountain....”.

Melica imperfecta Trin. MELIC-GRASS. “sandy flats, dunes...Point Lobos, Sunset...Lake

Merced”. Also above Baker Beach.

Phalaris californica Hook. & Arn. [CALIFORNIA CANARY-GRASS]. “along streams in

the sand hills and coastal bluffs: Point Lobos...Laurel Hill cemetery...”. Probably extirpated

in dunes here.

Poa douglasii Nees. DUNE BLUEGRASS. “Occasionally forming colonies on dunes and

sandy flats: ‘sand near the sea....Point Lobos, Richmond district, Sunset district, Lone

Mountain.”. Occurs today at Baker Beach, Fort Funston, Sunset Heights, dunes above Balboa

and Great Highway.

Poa unilateralis Vasey. OCEAN-BLUFF BLUEGRASS. “dunes, Sunset district.” Possibly

extirpated on dunes.

Vulpia microstachys (Nutt.) Benth. var. microstachys (Festuca megalura Nutt.) [ANNUAL

FESCUE] “common on...dunes...near Bakers Beach...Presidio...dunes, Sunset District”.

Vulpia octoflora var. hirtella (Walter) Rydb. (Festuca octoflora var. hirtella Piper) 

[ANNUAL FESCUE] “sandy slopes above Bakers Beach...”.

TYPHACEAE

Typha dominguensis Pers. SOUTHERN CATTAIL. “wet and marshy places...dunes, Sunset

district”. This collection undoubtedly refers to perennially moist dune slack.
Appendix I - 18 

Literature Cited

Best, C., J. T. Howell, W. and I. Knight, and M. Wells. 1996. A Flora of Sonoma

County. California Native Plant Society. 347 pp.

Bolander, H. N. 1863. Enumeration of shrubs and trees growing in the vicinity of the

mouth of San Francisco Bay. California Academy of Sciences Proceedings

2:78-84.

Brandegee, K. 1892. Catalogue of the flowering plants and ferns growing

spontaneously in the city of San Francisco. Zoe 2:334-383.

Clary, R. H. 1980. The Making of Golden Gate Park. California Living Books, San

Francisco Examiner Division of the Hearst Corporation, San Francisco,

California.

Hickman, J. C. (ed.). 1993. The Jepson Manual: Higher Plants of California. 

University of California Press. Berkeley, California.

Howell, J. T. 1949. Marin Flora. University of California Press; Berkeley and Los

Angeles, California. 319 pp.

Howell, J. T., P. H. Raven, and P. R. Rubtzoff. 1958. A Flora of San Francisco,

California. The Wasmann Journal of Biology 16:1-157.

Kaufeldt, C. E. 1954. Observations of San Francisco’s last major sand dune, with

notes on its flora and insect fauna. The Wasmann Journal of Biology 12:293-

336.

McClintock, E., P. Reeberg, and W. Knight. 1990. A Flora of the San Bruno

Mountains. California Native Plant Society Special Publication number 8.

Sacramento, California.

McLaren, J. 1924. Gardening in California: Landscape and Flower. A.M. Robinson 

Publishers; San Francisco, California.

Munz, P. A. 1959. A California Flora. University of California Press. 

Skinner, M. W. and B. M. Pavlik. 1994. California Native Plant Society’s Inventory

of Rare and Endangered Vascular Plants of California, 5th edition. CNPS

Publications, Sacramento, California. 338 pp. 

Thomas, J. H. 1961. Flora of the Santa Cruz Mountains of California. Stanford

University Press; Stanford, California. 
Appendix I - 19 

Wood, M. 1996. Rare and Endangered Plants of San Francisco’s Wild and Scenic

Places, 1st edition. California Native Plant Society, Yerba Buena Chapter.

Personal Communications

Holloran, P. 1998. California Native Plant Society, Yerba Buena Chapter, San

Francisco, California.

Thomas, T. 1997. Golden Gate National Recreation Area, National Park Service, San

Francisco, California.
Appendix II - 1

APPENDIX II. Native Vascular Plant Species of Serpentine Outcrops 

of the Northern San Francisco Peninsula

The following list of vascular plants occurring on serpentine outcrops in San Francisco

is compiled from Howell et al. 1958 and McCarten 1986, based on explicit references

to species growing on serpentine soils. Species included parenthetically [ ] indicate

either author’s observations, or species that are not explicitly cited by Howell et al.

(1958) as occurring on serpentine (e.g. some species recorded at Bayview Hills), but

are interpreted as likely occurrences on serpentine due to generally observed affinity

for serpentine sites, and occurrence at localities with known serpentine inclusions (and

other records of typical serpentine species). The “serpentine” of botanists may include

greenstone (identified as KJg on U.S. Geologic Survey maps of San Francisco),

volcanic rocks that sometimes resemble the grayish blue-green serpentine rocks, but

have less extreme forms of the distinctive chemistry of serpentine. Serpentine

occurrences are indicated in U.S.G.S. maps as “sp” and “sph”; they may occur as

inclusions in “Ks” (sheared rocks).

Nomenclature follows Hickman 1993, with synonymy provided from Howell et al.

1958. English names are compiled from primarily Best et al. 1996 and Hickman 1993. 

English names given in brackets are artificial English translations provided when

conventional common names are lacking.

The list does not cover mosses and lichens, which may occur in local abundance on

bedrock outcrops. Species marked with an asterisk * are reported to occur often on

serpentine, at least in the San Francisco flora. Parenthetic question marks (?) indicate

either some degree of taxonomic uncertainty, or less certainty of occurrence on

serpentine. Historic collection localities that include references to serpentine soils

include Potrero Hills, Laurel Hill, Masonic Cemetery, Presidio, and Hunters Point.

 
Appendix II - 2

- Pteridophytes -

(Ferns and fern-like plants)

Polypodiaceae

Polypodium californicum

CALIFORNIA POLYPODY FERN

Polypodium scouleri

LEATHERY POLYPODY FERN

Pteridium aqualinum

BRACKEN FERN

Pityogrammma triangularis

GOLDBACK FERN

- Angiosperms or Anthophytes -

(Flowering plants)

DICOTS

(broad-leaf flowering plants)

Apiaceae

Daucus pusillus

RATTLESNAKE WEED

Lomatium caruifolium var. caruifolium

[CARAWAY-LEAF LOMATIUM]

Lomatium dasycarpum ssp. dasycarpum*

[LOMATIUM]

Perideridia kelloggii*

[KELLOGG’S SQUAWROOT]

Sanicula arctopoides

YELLOW MATS

Sanicula bipinnatifida*

PURPLE SANICLE

Asteraceae

Achillea millefolium

YARROW

Agoseris apargioides

COAST DANDELION

Artemisia californica

CALIFORNIA SAGEBRUSH

[Aster chilensis]

[CHILEAN ASTER]

Baccharis pilularis

COYOTE-BRUSH

Cirsium andrewsii

FRANCISCAN THISTLE

Cirsium quercetorum

BROWNIE THISTLE

Erigeron glaucus

SEASIDE DAISY

[Eriophyllum staechadifolium]

LIZARD-TAIL

Filago californica

[CALIFORNIA FILAGO]

Gnaphalium microcephalum

[SMALL-HEADED CUDWEED]

Gnaphalium purpureum

[PURPLE CUDWEED]

Grindelia hirsutula var. maritima (G.

maritima)

SAN FRANCISCO GUMPLANT

Hesperevax sparsiflora (Evax sparsiflora)

[FEW-FLOWERED EVAX]

[Helianthella castanea]

DIABLO HELIANTHELLA

Lasthenia californica (Baeria chrysostoma)

CALIFORNIA GOLDFIELDS

Layia platyglossa

TIDY-TIPS

Lessingia filaginifolia var. californica

CALIFORNIA-ASTER

Microseris bigelovii

[BIGELOW’S MICROSERIS]

Microseris douglasii

[DOUGLAS’ MICROSERIS]

Psilocarphus tenellus

WOOLY-MARBLES
Appendix II - 3

Solidago spathulata

COAST GOLDENROD

[Wyethia angustifolia] (?)

MULE-EARS

Uropappus linearifolius (Microseris

linearifolia)

SILVER-PUFFS

Boraginaceae

[Cryptantha flaccida]

CRYPTANTHA

(?) Plagiobothrys reticulatus var. rossorianum

(P. diffusus)

GREENE’S POPCORNFLOWER**

Brassicaceae

[Arabis blepharophylla]

COAST ROCKCRESS

Lepidium nitidum

SHINING PEPPERCRESS

Caryophyllaceae

Spergularia macrotheca

STICKY SAND-SPURREY

Minuartina pusilla

[LITTLE SANDWORT]

Convolvulaceae

[Calystegia purpurata ssp.

purpurata](Convolvulus occidentalis var.

purpuratus)

MORNING-GLORY

Calystegia subacaulis

HILL MORNING-GLORY

Crassulaceae

Dudleya farinosa

[COAST DUDLEYA]

Ericaceae

Arctostaphylos hookeri ssp. franciscana*

(A. franciscana)

FRANCISCAN MANZANITA

Arctostaphylos hookeri ssp. ravenii*

(A. franciscana misappl.)

RAVEN’S MANZANITA

Fabaceae

Astragalus gambellianus*

GAMBELL’S MILKVETCH

Lotus wrangelianus (L. subpinnatus)

CALF LOTUS

Lupinus nanus

[DWARF LUPINE]

Lupinus variicolor

[COAST LUPINE]

Lupinus bicolor`

MINIATURE LUPINE

Trifolium depauperatum var. amplectens (T.

amplectens)

PUFFY CLOVER, BLADDER 

CLOVER

Trifolium fucatum (T. flavum)

BULL CLOVER

Trifolium gracilentum var. gracilentum

PINPOINT CLOVER

Trifolium macraei

CHILE CLOVER

Trifolium microdon

SQUARE-HEAD CLOVER

Trifolium microcephalum

MAIDEN CLOVER

Gentianaceae

Centaurium muehlenbergii

CANCHALAGUA

Hydrophyllaceae

Phacelia californica

[CALIFORNIA PHACELIA]

Lamiaceae

Stachys ajugoides var. rigida

(S. rigida ssp. quercetorum)

HEDGE-NETTLE

Linaceae

[Hesperolinon californicum]

[CALIFORNIA DWARF-FLAX]

Hesperolinon congestum*

MARIN DWARF-FLAX

Onagraceae

Clarkia franciscana*

PRESIDIO CLARKIA
Appendix II - 4

Orobanchaceae

[Orobanche californica ssp. californica]

CALIFORNIA BROOM-RAPE

Orobanche fasciculata

CLUSTERED BROOM-RAPE

Papaveraceae

Eschscholzia californica

CALIFORNIA POPPY

Platystemon californicus

CREAM-CUPS

Plantaginaceae

Plantago erecta

[ERECT PLANTAIN]

[Plantago maritima]

SEASIDE PLANTAIN

Plumbaginaceae

[Armeria maritima ssp. californica]

CALIFORNIA SEA-PINK

Polemoniaceae

Gilia clivorum

GILIA

Polygonaceae

Eriogonum latifolium

COAST BUCKWHEAT

Eriogonum nudum (?)

NUDE BUCKWHEAT

Pterostegia drymarioides

PTEROSTEGIA

Portulacaceae

Calandrinia ciliata

RED MAIDS

Claytonia exigua ssp. exigua (Montia

spathulata)

CLAYTONIA

Claytonia perfoliata

MINER’S-LETTUCE

Ranunculaceae

Ranunculus californicus

CALIFORNIA BUTTERCUP

Rhamnaceae

Ceanothus thyrsiflorus

BLUE-BLOSSOM

Rosaceae

Aphanes occidentalis (Alchemilla occidentalis)

WESTERN LADY’S MANTLE

[Fragaria chiloensis]

BEACH STRAWBERRY

[Oemleria cerasiformis]

OSO BERRY

[Heteromeles arbutifolia]

TOYON

Rubiaceae

Galium porrigens (G. nuttallii misappl.)

CLIMBING BEDSTRAW

Scrophulariaceae

Castilleja affinis ssp. affinis

COAST INDIAN-PAINTBRUSH 

Castilleja subinclusa ssp. franciscana (C.

franciscana)

FRANCISCAN INDIANPAINTBRUSH

Collinsia multicolor (C. franciscana)

FRANCISCAN CHINESEHOUSES

[Mimulus aurantiacus]

STICKY MONKEYFLOWER

Mimulus guttatus

LARGE MONKEYFLOWER

Triphysaria floribunda (Orthocarpus

floribundus)*

SAN FRANCISCO OWL’SCLOVER

Scrophularia californica

BEE-PLANT

Violaceae

Viola pedunculata

WILD PANSY
Appendix II - 5

- MONOCOTS -

 (Flowering plants with single 

 seedling-leaves or cotyledons)

Cyperaceae

Carex densa

DENSE SEDGE

Carex gracilior

SLENDER SEDGE

Juncaceae

Juncus bufonius

TOAD RUSH

[J. occidentalis]

[WESTERN RUSH]

Iridaceae

[Iris douglasiana]

DOUGLAS IRIS

[Sisyrinchium bellum]

BLUE-EYED GRASS

Liliaceae

Allium dichlamydium

COAST WILD ONION

Brodiaea terrestris ssp. terrestris

GROUND BRODIAEA

Chlorogalum pomeridianum

SOAP-PLANT

Dichelostemma capitatum (Brodiaea pulchella)

BLUE-DICKS

Muilla maritima

MUILLA

Triteleia laxa (Brodiaea laxa)

ITHURIEL’S SPEAR

Zigadenus fremontii

STAR-LILY

Poaceae

Agrostis exarata

SPIKE BENT-GRASS

Agrostis pallens (A. diegoensis)

THIN BENT, DUNE BENT

Agrostis microphylla

[SMALL-LEAF BENT-GRASS]

Bromus carinatus var. carinatus

CALIFORNIA BROME

Danthonia californica

CALIFORNIA OATGRASS

Deschampsia cespitosa ssp. holciformis

PACIFIC HAIRGRASS

Deschampsia danthonioides

ANNUAL HAIRGRASS

Elymus glaucus ssp. virescens (Elymus

virescens)

[GREEN WILDRYE]

[Elymus glaucus ssp. glaucus]

BLUE WILDRYE

Festuca idahoensis

IDAHO FESCUE, BLUE 

BUNCHGRASS

[Festuca rubra]

RED FESCUE

Hordeum brachyantherum

MEADOW BARLEY

Hordeum jubatum (Sitanion jubatum)

SQUIRRELTAIL BARLEY

Koehleria macrantha (K. gracilis)

JUNEGRASS

Vulpia microstachys var. pauciflora (Festuca

reflexa, F. pacifica)

[PACIFIC FESCUE]

Melica californica

CALIFORNIA MELIC

Melica torreyana*

TORREY’S MELIC

Nasella pulchra (Stipa pulchra)*

PURPLE NEEDLEGRASS

Poa secunda ssp. secunda (P. scabrella)

PACIFIC BLUEGRASS

Poa unilateralis

OCEAN-BLUFF BLUEGRASS

** Appears on McCarten’s list of Presidio

serpentine species. No reference to occurrence

in serpentine in Howell et al. 1958, or implicit

in collection locality.
Appendix II - 6

Literature Cited

Best, C., J. T. Howell, W. and I. Knight, and M. Wells. 1996. A Flora of Sonoma

County. California Native Plant Society. Sacramento, California. 347 pp.

Hickman, J. C. (ed.). 1993. The Jepson Manual: Higher Plants of California. 

University of California Press. Berkeley, California.

Howell, J. T., P. H. Raven, and P. R. Rubtzoff. 1958. A Flora of San Francisco,

California. The Wasmann Journal of Biology 16:1-157.

McCarten 1986. Serpentines of the San Francisco Bay Region: vegetation, floristics,

distribution, and soils. Unpublished report prepared for the California

Department of Fish and Game (revised 1993), Sacramento, California.
Appendix III - 1

APPENDIX III. Invasive Nonnative Plants That May Affect Recovery of 

San Francisco Lessingia and Raven’s Manzanita

Conicosia pugioniformis (L.) N.E. Br. [YELLOW ICE-PLANT]. This species is a

short-lived succulent perennial herb with a single tough taproot and prostrate branches

that spread radially up to a few meters. Unlike iceplant (Carpobrotus edulis and

hybrids), this species does not spread clonally by rooting prostrate stems of indefinite

growth. This species spreads readily by seed in coastal dunes, and is becoming

aggressively invasive around Baker Beach and Lobos dunes. It is native to South

Africa, but is invasive in numerous dunes systems of the central California coast.

Carpobrotus edulis (L.) N.E. Br. and hybrids with Carpobrotus chilensis (L.) N.E. Br.

ICEPLANT, HOTTENTOT-FIG. This species is the familiar iceplant of roadside

plantings, a creeping soft-wooded succulent shrub capable of indefinite lateral clonal

spread. Stems can root in contact with moist soil or sand. Carpobrotus edulis also

spreads efficiently by seed dispersed by birds and mammals (D’Antonio 1990). 

Intermediate hybrids and introgressants with the less aggressive Carpobrotus chilense

are common (Albert et al. 1997, Gallagher et al. 1997). It is a dominant species on

coastal dunes and sandy bluffs, covering tens of hectares of Fort Funston dunes in

nearly pure stands. Iceplant also occurs in the Presidio, where it covers both

serpentine and weak sandstone bluffs. Iceplant is shade-tolerant enough to form an

understory beneath conifers and eucalypts planted on remnant dunes of the Presidio. 

This native of South Africa can be effectively removed by labor-intensive manual

methods. The herbicide glyphosate is highly effective at eradicating it.

Cortaderia jubata (Lemoine) Stapf JUBATA GRASS, PAMPAS GRASS. The

familiar huge plumed fruiting culms of this large (to 3 meters or more) tussockforming grass are among the dominant features of coastal vegetation along the central

coast. This highly invasive species is distinguished from the ornamental pampas grass

(Cortaderia selloana) by the hairy sheaths on its flowering culms. It is tolerant of

most serpentine soils, as well as other Franciscan rocks, marine terrace deposits, and

sandstones. Jubata grass rapidly invades disturbed soils of slumps on coastal bluffs of

the Presidio, particularly where seeps occur. It is native to western montane South

America. This highly persistent invasive species is most easily controlled only at the

seedling/juvenile stage. Removal of mature tussocks by manual methods is effective,

but labor-intensive. It can also be controlled effectively by the herbicide glyphosate,

but only if coverage is thorough, since individual shoots that are not exposed can

vegetatively regenerate independently of the parent plant.

Bromus diandrus Roth RIPGUT BROME. This species and the Mediterranean

annual bromegrass B. hordeaceus readily invade coastal grassland, especially older

dunes with incipient soil development. It is particularly invasive in Lessingia

germanorum colonies in the Presidio. Other invasive annual exotic grasses include
Appendix III - 2

Avena fatua (wild oat), and Briza spp. (Briza minor, Brizqamaxima; rattlesnakegrasses). They can be controlled by labor-intensive manual removal, but they reinvade

readily unless extensive areas are cleared completely for multiple years. In managed

natural areas with infrequent native grasses, they can be controlled with grass-specific

herbicides.

Cupressus macrocarpa Gordon MONTEREY CYPRESS. This ornamental conifer is

a native tree of the Monterey peninsula. It was extensively planted in San Francisco,

where it has naturalized on coastal bluffs. Large groves cover remnant dunes at the

Presidio and Fort Funston. It also is tolerant of many serpentine soils. It

vegetationally regenerates from seed and invades coastal scrub vegetation on the San

Francisco peninsula, creating a dense shaded understory and thick acidic leaf litter

layer.

Ehrharta erecta Lam. UPRIGHT VELDTGRASS. This South African perennial

grass is a relatively recent invader. It has a tufted to mat-forming growth habit, and

forms an extremely tough sod. It is particularly invasive in bedrock crevices and

sidewalk cracks. It invades exposed stable coastal dunes in full sun, but is most

abundant and dominant on dunes shaded by Monterey cypress, and on relatively mesic

north slopes. It spreads aggressively by seed over relatively long distances, and

spreads locally also by clonal growth. Upright veldtgrass is locally abundant in parts

of the Presidio, Sunset, and Richmond districts of San Francisco, and is established on

southwest Farallon Island (San Francisco). Tenacious fibrous roots make manual

removed very difficult. It is likely to spread throughout bedrock and dune habitats

throughout San Francisco and other parts of the coast. It is sensitive to the herbicide

glyphosate.

Eucalyptus globulus Labill. BLUE GUM. This large evergreen Australian tree was

planted extensively on San Francisco dunes as well as on other substrates. It resprouts

readily after cutting. Its evergreen leaves are resistant to decomposition and produce

thick leaf litter that inhibits seedling regeneration of many native plant species. 

Seedlings establish spontaneously almost wherever mature trees occur in San

Francisco.

Genista monspessulana (L.) L. Johnson. FRENCH BROOM. This evergreen,

drought-tolerant, nitrogen-fixing shrub with sweetly scented and showy yellow

flowers has spread from cultivation. It produces copious seeds that can persist for

many years in the soil. Broom aggressively invades excavated banks, landslides,

artificial fill, and rocky open slopes; it seldom is invasive on dune sand. It regenerates

readily from seed and from resprouts of trunks. Manual removal is difficult, and

seedlings regenerate cleared stands for years after mature plants have been removed.

Pinus radiata D. Don MONTEREY PINE. Like Monterey cypress, this ornamental

conifer is a native tree of the Monterey peninsula. It is particularly well adapted to
Appendix III - 3

sandy soils, but also colonizes serpentine coastal bluffs, spreading by seed readily. 

Large groves also were planted in the Presidio on relict dunes because of their

tolerance to wind and salt spray. Plants do not resprout after cutting.

Literature Cited

Albert, M. E., C. M. D’Antonio, and K. A. Schierenbeck. 1997. Hybridization and

introgression in Carpobrotus spp. (Aizoaceae) in California. I. Morphological

evidence. American Journal of Botany 84:896-904.

D’Antonio, C. M. 1990. Seed production and dispersal in the non-native, invasive

succulent Carpobrotus edulis (Aizoaceae) in coastal strand communities.

Journal of Applied Ecology 27: 693-702.

Gallagher, K. G., K. A. Schierenbeck, and C. M. D’Antonio. 1997. Hybridization and

introgression in Carpobrotus spp. (Aizoaceae) in California. II. Allozyme

evidence. American Journal of Botany 84:905-911.
Appendix IV - 1

APPENDIX IV. Glossary of Technical Terms

achene: a seed-like hard-coated fruit containing one seed, and functioning as one

seed.

backcross: sexual reproduction between a hybrid organism and an individual from

one of its parent populations.

basal rosette: a plant growth form or stage with an unelongated stem near ground

level, and with a spiral circle of leaves.

blowout: a wind-eroded, mobile, mostly unvegetated portion of a dune, often forming

a depression.

building: The terms “buildings”, “infrastructure”, and “structures” include the

earthworks around the gun batteries, and other similar artificial topography that was

“built” to fulfill the organic functions of the military installations at the Presidio, and

any prehistoric features as well (such as hunting blinds, middens, or any mounds,

holes, or trenches associated with campsites, burial grounds or other human use sites).

CAS: abbreviation for herbarium specimens of the California Academy of Sciences,

Golden Gate Park, San Francisco.

Colma formation, Colma deposits: ancient deposits of mostly fine sand, variably

cemented, from “fossil” lagoon and dune environments; now found well above

modern sea level elevations both on the coast and inland along the northern San

Francisco peninsula, resembling yellowish modern dune sand.

colonize, colonization: establishment of a species’ local population or colony in a

given area.

competition: in plants, the interference with growth or reproduction in one plant that

is related to the exploitation or capture of needed resources (such as light, water,

space, nutrients, pollinators, shelter) by another plant. 
Appendix IV - 2

DS: abbreviation for herbarium specimens of the Dudley/Stanford collection, housed

at the California Academy of Sciences, San Francisco.

decumbent: growth habit with stems nearly prostrate, but ascending near tips.

disturbance: in the context of plant ecology, physical displacement of substrate or

physical removal of biomass in a given area.

dune, dune sheet: an unconsolidated deposit of wind-blown sand in modern

environments. A dune sheet is the mantle of sand covering underlying substrate

formed by a wide, continuous deposit of dunes.

endemic: narrowly and uniquely restricted to a single geographic area.

eolian: related to or caused by wind, such as erosion or deposition.

extirpate: killing of a population; local extinction.

facultative: an organism that does not require, but often behaves, tolerates or thrives

in, a particular condition or relationship, such as a facultative parasite (does not

require a host, but readily can behave as a parasite) or facultative salt-tolerant plant

(does not require salt, but can adapt to saline conditions).

fetch: the reach of open area across which wind may blow without obstruction.

floret: a small component flower within a larger structure resembling a single

blossom that is a composite of many small florets.

forb: an herbaceous plant.

founder(s): the pioneering individual or individuals that colonize a new site or

establish a new population; earliest ancestors of a distinct population.
Appendix IV - 3

Golden Gate National Recreation Area: a unit of the National Park Service, U.S.

Department of the Interior, located between Tomales Bay, Marin County, and north

coastal San Mateo County.

greenstone: with serpentine, mafic and ultramafic igneous rocks, derived from altered

minerals of deep magmas rich in heavy metals.

Holocene: a geological epoch dating back from modern times to the end of the

Pleistocene epoch when the last glaciers of the continent retreated and sea level rose

rapidly, about 13,000 years ago.

hummocky: having numerous small mounds

hybrid: the progeny of sexual reproduction between organisms from distinct and

separate lineages, such as different species, varieties, forms, or populations.

inbreeding, inbred: sexual reproduction between closely related individuals.

interspecific: between species (such as interspecific competition, interspecific

hybridization, interspecific differences).

introgression, introgressant: The result of backcrossing hybrid individuals to a

parent population, such that hybrids are assimilated, and their descendants fall largely

within the range of variation of the one ancestral parent population; such individuals

are called introgressants.

invasive: pertaining to species that rapidly establish large populations, becoming

dominant, often displacing pre-existing species; “aggressive” or “weedy” species.

involucre: a structure on flower heads composed of numerous scale-like or leaf-like

appendages (bracts) united to enclose unexpanded flower heads in bud-like coverings.

Merced: Ancient (Pleistocene) deposits of shallow marine or lagoon sediments

ranging from clay to sand, mostly along the modern coastline and raised above modern

sea level; named after the type locality at Lake Merced (Fort Funston) bluffs.
Appendix IV - 4

moist-chilling: a horticultural treatment of seeds to stimulate germination; moist,

imbibed seed are held at very cool above-freezing temperatures for many weeks;

“stratification” is an older term.

nonnative: exotic, or not indigenous to an area; introduced artificially or very

recently established without precedent in ecological time-scales.

outcross: opposite of inbreeding; sexual reproduction between individuals within a

species but from different lineages (families).

phyllary: a type of bract (see bract, above) in flower heads of aster (daisy) family

species.

Pleistocene: a geologic epoch characterized by many cycles of glaciation (advance

and retreat of continental glaciers, or ice-ages) and corresponding rise and fall of sealevel; ended with the last glaciation and the beginning of the Holocene.

self-sustaining: the maintenance of a biological resource which insures replacement

in perpetuity of individuals in the population through natural reproduction.

serpentine, serpentinite: mineral composition extremely deficient in key plant

nutrients such as calcium and magnesium, and rich in potentially toxic metals such as

cadmium, manganese, and nickel. Serpentinite is the rock itself.

significant net growth: Clones that are growing at a rate that is not significantly

lower (at the 95 percent confidence level) than the growth rates of healthy individuals

of a closely related species, the Franciscan manzanita (Arctostaphylos franciscana)

growing in interior sites on the San Francisco Peninsula.

slack: a relatively flat, low area within a dune system where surface elevations are

close to the permanent water table, and support either dune wetlands or wetland

ecotones.

soil seed bank: a population of dormant seed in the soil, functioning as a pool or

reservoir from which seedlings may be recruited. “Seed bank” is often used in
Appendix IV - 5

ecological contexts without the qualifying term “soil”. In plant conservation contexts,

the terms “banked seed” or “seed banks,” in contrast, refer to artificially stored seed.

succession: a developmental pathway of ecological change, typically involving shifts

in vegetation composition and structure as well as soil change; partly analogous with

orderly development of individual organisms, but more indeterminate and affected by

circumstantial or chance factors.

taxon (singular), taxa (plural): any unit of biological classification, such as variety,

subspecies, species, genus, etc. 

UC: abbreviation for herbarium specimens from the University of California

(Berkeley).
Appendix V - 1

APPENDIX V. Summary of Threats and Recommended Recovery Actions

LISTING

FACTOR

THREAT RECOVERY

CRITERIA

RECOVERY ACTION NUM

San Francisco Lessingia

A Urban development* 1(b) A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.3A Invasion of alien plants* 1(a)(i)(1-5)

1(a)(ii)(1)

A.1.1.1, A.1.1.2, A.1.1.3, A.1.2.1, A.1A.2.1, A.2.2, A.2.4, A.3, A.4.3

A Stabilization of sand in dunes* 1(a)

1(b)

A.1.1.1, A.1.1.2, A.1.1.3, A.2.1, A.2.2A.4.8

A Sand quarrying and bulldozing* 1(a) A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.3

A, E Damage by pedestrians, bicycles,

and off-road vehicles*

1(a) A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.3

B Potential future collection pressure* N/A A.3

D Limited protection under California

State law*

N/A Beyond scope of recovery plan

D Conflicts in land management

priorities*

1(a)1(b)

A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.3

E Garbage dumping* 1(a)

1(b)

A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.3
LISTING

FACTOR

THREAT RECOVERY

CRITERIA

RECOVERY ACTION NUM

Appendix V - 2

E Fertilizer-contaminated runoff* 1(a)

1(b)

A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.3

E Habitat fragmentation* 1(a)

1(b)

A.1.1.1, A.1.2.1, A.1.2.2, A.1.2.3, A.2A.4.1, A.4.2, A.4.4, A.4.7

E Genetic and demographic threats

from small population size

1(a)1(b)

A.1.1.1, A.1.1.2, A.1.2.1, A.1.2.2, A.1A.2.4, A.4.1, A.4.2, A.4.4, A.4.7, A.5.

Raven’s Manzanita

A Urban development* 2(a)(i) B.2.7, B.3.2A Competition from other plants* 2(a)(i)

2(b)(ii)

B.1.1, B.1.2, B.1.3, B.2.2, B.2.4, B.2.8B.4. B.5.4, B.5.6, B.6.3

A, E Vulnerability to management errors

due to small population size*

2(a)(i)2(a)(ii)2(a)(iii)2(b)(i)2(b)(ii)2(b)(iii)

B.1.1, B.1.2, B.1.3, B.2.1, B.2.3, B.2.5B.3.3, B.3.4, B.3.6, B.3.7, B.4, B.6.1, 

B Scientific collection pressure* 2(b)(iii) B.4, B.6.3

C Fungal twig blight 2(b)(iii) B.2.1C Herbivory by tussock moth

caterpillars

2(b)(iii) B.2.1
LISTING

FACTOR

THREAT RECOVERY

CRITERIA

RECOVERY ACTION NUM

Appendix V - 3

D Limited protection under California

State Law*

N/A Beyond scope of recovery plan

D Transfer to Presidio Trust or private

ownership may increasedevelopment pressure

2(a)(i) B.2.7

E Possible lack of pollinators* 2(a)(iv) B.1.2E Lack of genetic diversity 2(a)(ii)

2(a)(iv)

B.5.1, B.5.2, B.5.5, B.5.7, B.6.1

E Low reproductive success 2(a)(iv)

2(b)(i)

B.1.2, B.2.1, B.5.2, B.5.3, B.5.4, B.5.5

E Possible hybridization with other

manzanitas

2(a)(iv) B.4, B.5.3, B.5.5, B.5.7, B.6.1, B.6.3

Listing Factors: A. The Present or Threatened Destruction, Modification, or Curtailment Of Its Habitat or Range 

B. Overutilization for Commercial, Recreational, Scientific, Educational Purposes (not a factor)C. Disease or Predation D. The Inadequacy of Existing Regulatory Mechanisms E. Other Natural or Manmade Factors Affecting Its Continued Existence

* Identified as a threat in the final listing rule.
Appendix VI - 1

APPENDIX VI. Agency and Public Comments on the Draft Recovery Plan for

the Coastal Plants of the Northern San Francisco Peninsula

A. Summary of Agency and Public Comment on the Draft Recovery Plan for

Coastal Plants of the Northern San Francisco Peninsula

In December 2001, we released the Draft Recovery Plan for the Coastal Plants of the

Northern San Francisco Peninsula for a 90-day public comment period, which closed

on March 4, 2002 (U.S. Fish and Wildlife Service 2001). This comment period was

extended for an additional 60 days to close on May 6, 2002 (U.S. Fish and Wildlife

Service 2002a) and was then reopened for another 60 days beginning in July 2002 to

close on September 9, 2002 (U.S. Fish and Wildlife Service 2002b). During the

comment period we received comments from the public, government agencies, and

non-governmental organizations. We requested peer review from the following

individuals:

Dr. Thomas Parker Department of Biology, San Francisco State University

Dr. Norman Ellstrand Department of Botany, Plant Sciences & Program in Genetics,

University of California - Riverside

Ms. Lisa Wayne City of San Francisco, Department of Recreation and Parks

Ms. Andrea Pickart Humboldt Bay National Wildlife Refuge, U.S. Fish and Wildlife

Service

Dr. Kristina Schierebeck Department of Biology, California State University -

Chico

Ms. Crima Pogge Expert in the flora of San Francisco

Dr. Bruce Pavlik Department of Biology, Mills College, Oakland, California

Dr. Donald Mahoney Strybing Arboretum and Botanical Gardens, Golden Gate Park,

San Francisco, California

Dr. Eric Knapp Western Ecological Research Center, U.S. Geological Survey

Dr. Jon E. Keeley Division of Environmental Biology, National Science

Foundation

Dr. Staci Markos Jepson Herbarium and Department of Biology, University of

California - Berkeley

Mr. Michael Vasey Department of Biology, San Francisco State University

Mr. Roman Gankin San Mateo County Parks and Recreation, Redwood City,

California
Appendix VI - 2

Ms. Holly Forbes Jepson Herbarium, University of California, Berkeley Botanical

Garden

Dr. Barbara Ertter Jepson Herbarium, University of California - Berkeley,

California

Dr. Thomas Daniel California Academy of Sciences, Golden Gate Park, San

Francisco, California

Comments were received from five of the peer reviewers.

This section summarizes general information about the comments we received during

the comment periods. A complete index of commenters (with affiliation if one was

identified) is available from our Sacramento Fish and Wildlife Office, 2800 Cottage

Way, Suite W-2605, Sacramento, California 95826-1888. All comment letters are

kept on file at this office. 

In addition to the peer review comments, we received 425 letters, postcards, faxes, and

electronic mail messages. The following summarizes the number received from

various categories of commenters:

Federal agencies . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

local governments .......................... 1

business and industry ........................ 1

environmental and conservation organizations . . . . 2

cultural conservancy groups .................. 5

individual citizens . . . . . . . . . . . . . . . . . . . . . . . . . . 413

academic and professional (includes peer reviewers) 5

Peer review comments on the draft plan were generally very supportive. One peer

reviewer asked for clarification on various aspects of the recovery plan. Their

comments included, for example: “The Draft plan is a well-conceived, well-written,

and scientifically sound document that is a credit to your organization.,” “. . .a strength

of the plan is the focus on process–both the underlying ecological processes that led to

habitat creation, as well as processes to be restored.,” “. . .commend. . innovative

approach. . .,” “. . .an experimental program to breed variation back into [Raven’s

manzanita] is an excellent strategy. . .” Suggestions and comments also included: 
Appendix VI - 3

– the goal of restoring a sexually reproducing population of Raven's manzanita

must be done carefully and may not work, ....

– a recommendation for a scientific advisory panel to assist recovery,

– San Francisco lessingia found at Daly City is distinct from that found at the

Presidio and it may be advisable to maintain them as separate types in the

Presidio and Southern Recovery Units, and

– the ecosystem approach proposed is sound, appropriate, and the best way to

succeed.

Some of the 430 comment letters were the result of some individuals sending

comments more than once. Many letters were concerned with aspects of the plan as

they were described in news reports. The chief issue among such commenters was

their concern that the size of the area to be restored to habitat for San Francisco

lessingia was larger than needed, with an impact on the heritage landscape trees at the

Presidio. They cited a variety of reasons for minimizing the area of tree removal.

There were concerns raised about: (1) the possibility of restrictions on recreation at

the Presidio and Fort Funston; (2) the cost of managing a larger area for plants that

have needed intensive management effort in the past; (3) the ability to do the work on

the schedule described in the plan’s implementation schedule; (4) technical points

about the manner of restoration and the relative priority of some of the components of

the restoration; and (5) the possibility of difficulties reconciling the goals of the

recovery plan and the goals for the historic and esthetic values of the heritage

landscape.

There were also comments about several issues that are beyond the scope of the

recovery plan.

Information and comments not incorporated into the recovery plan were considered

and noted and may be useful in the future. Major comments that were not

incorporated, or that require clarification in addition to their incorporation, are

addressed below.
Appendix VI - 4

B. Summary of Comments and Our Responses

Comment: Many of the comments from individuals include one or both of two key

points:

1. We should not spend $17 million to cut down trees.

2. We should use habitat that is already suitable.

Response: The draft recovery plan’s total cost of about $17 million includes $4.5

million to buy habitat that is already suitable. The remaining $12.5 million of costs

estimated in the draft recovery plan are spread over the two listed species that are the

primary subject of the recovery plan; five other listed species that co-occur with them;

and 33 other sensitive species that are not listed, but co-occur in the same habitats and

face the same threats as San Francisco lessingia and Raven’s manzanita. The costs of

conservation and recovery actions are, thus, spread over 40 species, not just San

Francisco lessingia. The recovery actions also cover a wide range of activities in

addition to vegetation management and habitat acquisition, including public outreach

and education, research on the most efficient recovery methods, and planning efforts

with partners and stakeholders.

In addition, the plan’s costs extend over 30 years and at least 19 locations extending

from the Presidio south to Fort Funston and Daly City. The money will be spent at

multiple sites to manage the species and habitat through all of the numerous actions in

the recovery plan.

Comment: The recovery plan will result in premature removal of Wherry Housing,

with the risk of preventing timely economic self-sufficiency by the Presidio Trust.

Response: At the request of the Presidio Trust, we have adjusted the final recovery

plan to reflect the Trust’s current schedule for demolition of Wherry Housing with

one-third of the housing to be removed in 10 years, another one-third in 20 years and

the final one-third in 30 years to allow economic self-sufficiency of the Presidio Trust.

Comment: The National Park Service and the Presidio Trust, as well as many

individual commenters, had concerns about meeting the schedule of the recovery

actions in the recovery plan. One concern of the National Park Service is that
Appendix VI - 5

Congress may not approve the full amount requested by the National Park Service in

coming years to help fund recovery actions.

Response: It is important to keep in mind that we view the recovery plan as a living

document, not a regulatory mandate. The plan outlines actions we believe will lead to

the recovery of these species. We, the U.S. Fish and Wildlife Service, are committed

to working closely with other agencies, landowners and other stakeholders to make

sure these recovery actions, including the schedule, work for all parties.

Cost estimates for the recovery actions do not assume the National Park Service or

Presidio Trust would receive additional funding on the exact schedule of the recovery

plan. This is reflected in the disclaimer in the Preface to the recovery plan. Nor does

the recovery plan assume that the National Park Service or Presidio Trust would

provide 100 percent of the money spent to do the recovery actions, even the ones

where they will be playing a leading role. We provide funding to landowners, other

agencies and partners across the country to help with recovery actions, and we intend

to help fund recovery actions called for in this plan as funding is available.

Comment: Blue gum trees (Eucalyptus globulus) are a public nuisance due to falling

limbs, excessive shedding of bark and branches, treefall hazards, low wildlife value,

and fire hazard. San Francisco has enough of them. There are more than enough

Monterey cypress (Cupressus macrocarpa) and Monterey Pine (Pinus radiata), and

their local removal in exchange for native biological diversity is welcome.

Response: Public nuisance factors of trees, in and of themselves, are not the subject

of this recovery plan. But we appreciate your providing us with your viewpoint. 

Description of incidental public benefits of tree removal, such as those cited, will be

most appropriate for analysis in a comprehensive environmental assessment or impact

statement under the National Environmental Policy Act (NEPA). The National

Environmental Policy Act review would be triggered by plans for individual recovery 

projects to be done by the lead Federal land management agencies.

Comment: Many commenters had concerns about the restoration of dune vegetation

where there are trees, including:

1. the esthetic value of the trees for visitors (compared with dune scrub plants)

2. the stabilizing effect of the trees on the sand
Appendix VI - 6

3. the sheltering effect of the trees from north winds

4. the spiritual value of the trees

5. the value of trees for absorption of carbon dioxide and air pollution

6. the historic value of the trees as part of a recreational area for military personnel

during World War II

7. the apparent high cost of the tree removal

8. the endangered plants are not worth the cost in terms of trees and dollars

9. adverse effects on wildlife, especially birds. More wildlife will be displaced by

tree removal than drawn by native dune scrub habitat restoration

10. the perceived large increase in tree removal compared to earlier plans

11. the entire restoration area currently has trees and will be replaced over the entire

area with San Francisco lessingia as the only species of plants

12. The Presidio's trees in urban San Francisco have a symbolic value that transcends

their esthetics. Trees provide a symbolic link to forests and nature, and are

experienced as connection to benign nature by city dwellers. Urban forests are

part of the European and North American sensibility.

13. Research should be done to investigate which trees could function as windbreak

and would be compatible with the National Historic Preservation Act and the

conservation needs of the San Francisco lessingia.

Several other commenters said they were in favor of restoration of native habitat. 

While some were concerned about the potential for reduced recreational opportunities,

others saw a potential for increased recreational opportunities, such as by being able to

visit a habitat that has largely disappeared or been degraded.

Response: The recovery plan does not recommend replacing Presidio trees with San

Francisco lessingia. It recommends dune scrub habitat restoration in areas of dune

scrub, restored dune scrub, housing, and pavement. Landscape trees in the areas to be

restored would be replaced not with solid stands of San Francisco lessingia, but with a

variable, diverse coastal dune vegetation of which San Francisco lessingia is a small

part and which includes shrubs and trees such as arroyo willow (Salix lasiolepis) and

live-oak (evergreen oak trees of the genus Quercus). The coastal dune scrub would

not be identical to the coyote brush-dominated remnants of dune scrub that are left

over from historic grazing and military land uses. Restored stable dune vegetation is

predominantly composed of flowering shrubs and herbs, with relatively small patches

of sparse sandy areas. Crissy Field dunes, a highly popular site of high visitor use in
Appendix VI - 7

the Presidio, is composed mostly of stable dune vegetation that has both scenic and

esthetic appeal. Other scenic areas in California with similar stable dune vegetation

and high popular esthetic appeal include portions of Monterey Bay and Point Reyes.

The value of endangered plants is not limited to the individual economic utility of a

given species at any one time. The recommendations of this recovery plan do not

include replacement of low-diversity non-native tree stands with single-species stands

of endangered San Francisco lessingia. Arroyo willow, live oak, and other tree and

shrub species are represented in naturally occurring dune ecosystems. Existing dune

ecosystems can serve as sites to determine appropriate densities and composition for

management purposes, and allow for evaluation of function for windbreaks, sand

stabilization, and value for wildlife.

The Presidio Recovery Unit overlaps sections of the historic forest that contributes to

the National Historic Landmark status of the Presidio. Many of the non-native trees

that are of concern in providing for the recovery of the San Francisco lessingia are part

of that forest. The Presidio Vegetation Management Plan identifies the limits of the

forest, identifies specific management zones within the forest, and prescribes

management strategies within these zones. Adaptive management strategies will be

developed to provide optimal recovery conditions while attempting to retain the

overall historic character of the forest. In cooperation with the Presidio Trust and the

National Park Service, iterative strategies of thinning, limbing, increasing space

between trees, and even change out of species to lower growing but historically

consistent trees may be tested to increase ecosystem function, such as wind fetch. In

addition, buffers along the edges of the historic forest should be developed to prevent

weed invasion. The Presidio Vegetation Management Plan identifies most of the

southwest corner of the historic forest as a Special Management Zone in which all of

these management strategies are anticipated, and for which this recovery plan

recommends development of management plans involving implementing and testing

these strategies. Monitoring of the effectiveness of these strategies will be essential to

the recovery process.

Comment: San Francisco lessingia habitat should be restored elsewhere, or the

species should be established elsewhere, not in San Francisco. Recommended

alternative locations include Point Reyes, Monterey, Lanphere Dunes (near Arcata in

Humboldt County), and Ocean Beach.
Appendix VI - 8

Response: San Francisco lessingia is native only to the San Francisco peninsula. 

Because of the lowered chance of success for recovery and other potential

consequences, our policy is to restrict re-establishment and recovery work to be within

the historic range of a species. Introducing a species well outside its native range can

cause risk of invasive (weedy) behavior. For example, yellow bush lupine (Lupinus

arboreus), a modest native shrub of the central California coast, has become a highly

invasive, aggressive wildland weed where it was introduced in Northern California

and Oregon. Introducing species outside their native range may also cause genetic

threats to invaded ranges of other species.

The recovery plan acknowledges that San Francisco lessingia would be likely to

establish along the stable dune roadside of the Great Highway, along Ocean Beach, if

it were reintroduced to Sutro Heights dune remnants. This is not a recommended

strategy, however, because sea level rise may erode the Great Highway and force its

replacement, because the habitat there is highly degraded by imported fill and weeds,

and because the habitat is unstable due to the high degree of wind scour. 

Establishment there would make a valuable short-term contribution to the conservation

of the species, but is not a surrogate for natural, persistent habitat within a community

of native species maintained mostly by natural processes.

Comment: Many commenters commented that the species in the recovery plan were

not worth conserving, or were not worth foregoing other benefits. 

Response: Those comments are noted. The recovery of listed species is mandated by

the Endangered Species Act. The Endangered Species Act reflects the value that the

American people place upon the natural resources of the United States, and their

diversity. The Endangered Species Act directs us to conserve endangered and

threatened species and the ecosystems upon which they depend. 

Comment: Please explain the reasoning behind the acreage requirements identified in

the draft recovery plan regarding the specific acreages for the expansion of the reserve

areas.

Response: Utilizing a single minimum acreage based on population viability analysis

was not possible, because many of the San Francisco lessingia populations are not

fully natural, but instead have been seeded, eliminating the ability to analyze
Appendix VI - 9

population viability to its fullest. Therefore, the acreage requirements identified in the

draft recovery plan are based on an evaluation of each site. Our approach was to look

at the elevation and exposure of each site, and the number of acres that were feasible

to restore. We then used this information to evaluate how much restoration would be

necessary to ensure natural maintenance of ecosystem processes. This process yielded

the minimum reserve areas specified in the draft recovery plan (2 hectares [5 acres], 3

hectares [7 acres] and 44 hectares [110 acres] for Rob Hill, Public Health Services

Hospital, and Lobos-Wherry-Baker, respectively).

Comment: San Francisco lessingia's endangerment is an example of "Darwinian

survival of the fittest." Its endangerment indicates that it is not worthy of survival. 

Conservation measures promoting its survival and recovery are artificial contrivances.

Response: The threats to San Francisco lessingia are not natural, nor an indication of

the species' fitness. The replacement of nearly all habitat within its historic range, and

the decline of remnant habitat by introduced trees and weeds, are the primary reasons

for its endangerment. San Francisco lessingia has the ability to thrive and expand

where stable dune habitat is of suitable quality and quantity. Legally, Federal agencies

are obligated to utilize their authorities to conserve endangered species. Only under

special circumstances can a government panel make a recommendation to deliberately

allow the extinction of a species threatened by human actions.

Comment: The target of 5 percent non-native vegetation cover is shown to be

unrealistic by the monitoring data for Lobos Creek, where weeds have represented 15

percent of the cover. Further, the 15 percent weed cover has been maintained only by

intensive management by hand weeding. Therefore, it is recommended that we adopt

a flexible target for non-native vegetation cover based on actual monitoring data.

Response: The recovery areas outlined in this plan includes a larger area than Lobos

Creek and thus will allow for more efficient weed-control methods, in conjunction

with a more natural disturbance regime. In contrast, managing small preserves is

difficult because of the “edge effect.” In a small preserve, much of the overall area is

near the edge, and close to adjacent, non-preserve areas. For example, the rate of nonnative seed movement into a small preserve results in higher occurrence of seeds per

unit area than for a larger preserve. The entire area is also influenced by other factors,
Appendix VI - 10

such as the shading effect of forest. With a larger area, a greater part of the habitat is

relatively farther away from the effects outside the edge.

In the very successful dune scrub restorations of larger areas of habitat, such as

Lanphere Dunes near Arcata, it has been shown that keeping weed cover below the

threshold for explosive weed population growth is quite feasible. For these reasons,

we expect that the larger area of restored habitat will allow the smaller amounts of

invasive cover to be a manageable goal.

The recommendations for invasive cover are our best estimate of the levels needed to

minimize the need for large amounts of hand-weeding. The weed management

recommendations are based on the principles of weed management that call for prompt

identification of colonies of an invasive species that can be treated for eradication,

rather than treating entire areas for suppression. These principles are expected to be

applied to newly restored areas that begin with low or no invasive species cover, and

to areas where invasive species have been essentially eradicated. For the core

populations on larger areas of restored habitat, the intent is that most of the invasive

cover be kept from moving inside the outer margins of the restored dune scrub habitat. 

This will change the weed control need from suppression over the whole site to

suppression at the edges and removal of new colonists in the core area before they can

spread and require whole site treatment. Experience with new methods may prove that

somewhat higher or lower levels of invasive species as proportions of total cover and

plant cover may be the most efficient means to maintain dune scrub habitat, in which

case the new information would provide the basis for future management.

Comment: One commenter wanted us to consider deleting the statement that the

expansion of the Lobos Dunes population should precede restoration at any other

reserve.

Response: We have considered this suggestion, but have not adopted it because we

believe that the restoration at the Lobos Dunes population is a very high priority

action. It will be the largest population of San Francisco lessingia, expanding from an

existing population with its full range of genetic variation, and we feel it is essential to

the species recovery.
Appendix VI - 11

Comment: Please revise the statement on page 127 of the draft plan, “Incompatible

uses (planted trees or dense shrub cover, road construction, irrigation, fertilizer

application, construction of buildings, etc.) must be excluded from Recovery Unit in

perpetuity,” and consider replacing it with, “The Presidio Trust will identify

compatible uses and the appropriate management actions in the area and will consult

with us as required. If desired by both agencies, a programmatic agreement could

identify when further consultation is necessary for operational activities. Actions to

consider in this document include planted trees or dense shrub cover, road

construction, irrigation, fertilizer application, construction of buildings.”

Response: We are modifying the language to provide more flexibility in the

management of both the dune scrub restoration and surrounding landscape areas. In

both cases, it is suitable to use some supplemental watering to establish new

vegetation. However, we want to avoid providing water and fertilizer to the extent that

it would result in more weeds.

Comment: One comment suggested that other options besides wind be considered to

promote disturbance within San Francisco lessingia habitat.

Response: The recovery plan does provide for other options besides wind to promote

disturbance. Recovery Action A 4.5 recommends the experimental investigation of

the feasibility of rotational trail use as a vegetation management technique. Recovery

Action A 4.6 recommends conducting field experiments to determine the utility and

restoration feasibility of managing dune topography via human disturbance.

Comment: The draft recovery plan recommendations will reduce the esthetic and

recreational values of affected portions of the Presidio, and adversely modify local

climate. This would result in an appreciable decline in real estate values of adjacent

and neighboring private properties.

Response: The esthetic impacts of habitat restoration in the Presidio Recovery Unit

may not necessarily be adverse. There are regional indications that residences

adjacent to vegetated coastal dune scenic views command exceptionally high real

estate values. For example, the Monterey Bay Dune Colony, Marina Dunes

condominiums, Spanish Bay development (Monterey Bay), Ward Avenue homes

(Cleone, Mendocino), and Irish Beach homes (Mendocino) are all residences with
Appendix VI - 12

direct views of adjacent vegetated dune landscapes with sale values which exceed

comparable non-coastal homes in the immediate region.

The private residential lands neighboring the Presidio Recovery Unit roughly parallel

Lake Street, between 25th and 16th Avenues, due south of the Lobos Dunes site, and

the portion of the Seacliff neighborhood adjacent to Baker Beach, near the mouth of

Lobos Creek, southeast of the Presidio Recovery Unit. The dominant and prevailing

winds in the Golden Gate area are generally from the west, with less frequent winds

from the east and northeast. The sheltering effects of trees depends on tree height and

distance downwind, and on topography (i.e., slope). Additionally, the beneficial

effects of trees diminish rapidly on flat areas and are not significant even short

distances upslope above tree height. Therefore, the best available science leads us to

believe that climatic variation as a result of habitat restoration activities will be

minimal. If adverse downwind impacts of tree removal are substantial, effective

mitigation measures would need to be identified and designed as part of the National

Environmental Policy Act process.

Comment: The National Park Service recommended we enter into a Participation

Plan with affected parties. They further stated that there was no formal coordination

on developing alternatives for recovery implementation that address both the recovery

of the species as well as minimizing the social and economic effects of the recovery

action.

Response: In developing the draft recovery plan, we coordinated with natural

resources management staff at the Golden Gate National Recreation Area through emails, telephone conversations, and meetings. We hope to establish a partnership with

the National Park Service, Presidio Trust, and other stakeholders in carrying out

recovery actions. We have added an action to develop a Participation Plan that will

provide for the recovery of the species while minimizing social and economic impacts. 

We will involve all affected interests in the recovery plan implementation process

through the development of the participation plan. The participation plan should

ensure that a feasible strategy is developed for all affected interests while providing

realistic and timely recovery of the species. We recognize that no recovery plan can

succeed without the close cooperation of land managers and other partners.
Appendix VI - 13

Comment: There was inadequate public (stakeholder) participation in the drafting of

biological recommendations in the recovery plan.

Response: While we have been criticized for not working closely enough with the

Presidio Trust and the National Park Service on preparation of the draft plan, we have

revised the final plan to incorporate information, comments, and recommendations

provided by the Presidio Trust and National Park Service and have included additional

actions for development of participation plans with our partners and stakeholders.

We would also note that the public comment period on the draft plan was re-opened

twice, drew more than 400 comment letters, and included numerous public meetings

that were attended by up to 45 people each. The drafting of the recovery plan

overlapped with the preparation of the Presidio Vegetation Management Plan, which

also drew much public comment. Our staff followed the development of the Presidio

Vegetation Management Plan, and recognized the public concerns that were

expressed, particularly by neighbors. The indirect input of public comment in the

Presidio Vegetation Management Plan was fully considered during the later stages of

preparation of the recovery plan. All of the parties included in the Presidio Vegetation

Management Plan process were sent notice of our draft plan availability, comment

periods, and public meetings.

Comment: There was no public notice or inadequate public notice for public

meetings and publication of the draft recovery plan.

Response: We are always striving to do a better job involving the public in our

activities. In this instance, we sent 1,574 copies of the draft plan to affected or

interested parties. Copies also were distributed to local libraries. We had three

comment periods over a period of more than 10 months, from December 4, 2001,

through September 9, 2002. Each was announced by notices published in newspapers

and the Federal Register. In addition, we held six public meetings and sent

announcements to more than a thousand people who had indicated an interest in the

planning process. We also attended 12 homeowners association meetings.

Comment: Has the U.S. Fish and Wildlife Service coordinated with administrators of

other agencies?
Appendix VI - 14

Response: Yes. During the development of the draft recovery plan, we directly

solicited information from the various agencies and coordinated extensively with staff

from the National Park Service and the Presidio Trust. During the comment period we

directly requested comments from the agencies, and extended the comment period

specifically to give the National Park Service time to finish preparing their comments.

Comment: San Francisco lessingia is an ugly weed and unworthy of restoration

efforts and resources. Weeds are not wanted or needed in the Presidio.

Response: The common name "vinegarweed" has been erroneously misapplied to San

Francisco lessingia. There are no published floras, field guides, botany books, natural

history guides, or historic references to this term as a synonym for San Francisco

lessingia. None of the major research publications or taxonomic works on this genus

apply this common name. San Francisco lessingia does not have any of the essential

characters of a weed except for being found on partially bare ground. It is not invasive

or persistent, displaces no native or desirable plants of its own accord, and naturally

causes no economic harm. Weeds are not only found on bare ground, but often persist

in the plant community even when we try to get rid of them. Regarding esthetics, the

abundant lemon-yellow flowerheads, produced during a season with few other

flowers, provides colorful wildflower displays. It is associated with a coastal dune

plant assemblage that is widely photographed and painted by artists along the central

coast, and featured in many natural history books, photo calendars, travel books, etc.

The recovery plan objectives are for conservation of endangered plant species, not

weeds. The plan does not recommend addition of weeds, but recommends removal of

many weeds, which are invasive, non-native plant species that do ecological or

economic harm.

Comment: Restoration of sand dunes, and reintroduction of endangered species in

San Francisco is a futile and misguided attempt by a minority of native plant

enthusiasts/naturalists to "turn back the clock," an exercise of biological nostalgia

rather than restoration. Restoration of wildlands in a developed city is an affront to

the values of civilization, and suggests that the restoration agenda may next target

Golden Gate Park or other urban sites with great public value.
Appendix VI - 15

Response: The scale of restoration recommended in the recovery plan is limited

mostly to areas of San Francisco that are already parklands managed in part or whole

as native habitat with public access. One exception is Wherry Housing, which had

been previously programmed for demolition, prior to our recommendations about the

use of the site for recovery of San Francisco lessingia. The scale of tree removal in the

Presidio is limited to no more than 30 hectares (75 acres) over the next 30 years, with

every acre to be analyzed in site-specific plans subject to full environmental review,

most of which has already been recommended or considered for removal by the

Presidio Vegetation Management Plan. Other than Wherry Housing, the draft plan

does not consider replacement of urban development with restored habitat. Nor does

the draft plan recommend or consider converting large areas of horticulturally

developed parkland (lawns, golf greens, maintained ornamental plantings, and other

such artificial landscapes) to restored native habitat. We believe that a measured

approach to the restoration, as recommended by the National Park Service, the

Presidio Trust, and the California Native Plant Society, is the best way to coordinate

the change from housing to park landscape and avoid scheduling conflicts.

Comment: Restoration of native species by removing non-native vegetation is not

justifiable. Just because a species is not native does not mean that it is inferior in

value. "Native" is relative, depending on how far back in time we assess a biological

community. Discriminating against species, such as trees, because they are not strictly

"native" is arbitrary. For example: Monterey cypress is currently native only to the

Monterey peninsula in California, but was previously (geologic time) more

widespread.

Response: The basic purpose of the recovery plan is to recover the San Francisco

lessingia, Raven’s manzanita, and associated species. Restoration of native habitat is

important to that purpose. To that end, the recovery plan primarily focuses on publicly

owned lands for which there are already plans to restore native habitat. For example,

most of the area of the recovery units is included in such plans as the National Park

Service’s and Presidio Trust’s Presidio Vegetation Management Plan. 

The recovery plan proposes no general exclusion policy for non-native species; rather,

it focuses on the recovery units and those invasive species which degrade habitat

quality for the listed species within those units. Those non-native species which are

invasive and threaten the survival of an endangered species are the focus of this plan.
Appendix VI - 16

Comment: Several commenters were concerned that restoration of sand dunes or San

Francisco lessingia is incompatible with, or inappropriate for, an urban park setting

where recreational space is at a premium. They perceive habitat restoration as a

euphemism or rationalization for fencing off parks from public access. They see the

draft recovery plan as an example of the pattern. They perceive the draft recovery

plan's recommendations as unacceptably restrictive, and believe carrying them out will

exclude recreational uses of the Presidio, and contribute to the progressive exclusion

of recreational uses of other parts of the Golden Gate National Recreation Area (such

as Fort Funston), and, perhaps, city parks as well.

Response: The recovery plan is not a master land use plan that specifically addresses

recreational use policy for Fort Funston and the Presidio; nor does it address the

cumulative impacts of changes in management of public lands, such as parks, on

recreation in San Francisco as a whole. It addresses public access and recreation only

in relation to the conservation of endangered species. The recommended combination

of public access for hiking (to aid keeping the vegetation cover somewhat open by

trampling) and cyclic closure (to let native vegetation re-establish while moving the

hiking to other areas) as a compatible tool for vegetation management attempts to

reconcile potential conflicts between endangered plant conservation and existing

recreational uses, without compromising the integrity of recovery, within the scope of

the draft recovery plan.

Comment: It is not possible to restore an "ecosystem,” since scientific understanding

of ecosystems is incomplete. Restoration is arbitrary: selecting one point in the past to

restore over other past states is not justified. "Rare Plant ecosystems" are unique and

cannot be duplicated.

Response: We believe that ecosystem restoration is usually feasible if the site has not

been altered too much, if the underlying ecological processes can be restored, and the

key species are available. Coastal dune restoration and management in California has

a substantial track record. Lanphere Dunes (Arcata) is an outstanding example of

coastal dune restoration. Locally, the Crissy Field dune restoration in the Presidio has

successfully reintroduced the majority of local dune species, and integrated well with

pre-existing dune remnants. These were done not by controlling every aspect of the

ecosystem, which is impossible. Rather, they were achieved by restoring the basic
Appendix VI - 17

environmental and biological foundations, and allowing the living system to

regenerate itself.

We believe that our recommendation to restore dune habitat is based on sound current

scientific information. We recommend restoring the dune ecosystem, because that is

the ecosystem that has supported the San Francisco lessingia, and the best available

science indicates that restoration of dune habitat is essential for the recovery of that

species.

Many examples exist of plant communities that either have been spontaneously

restored, through succession after abandonment, or have been actively restored. Many

contemporary populations of rare and endangered plants occur on historically recent

sites following human land use changes. Most rare dune plants in California occur in

non-pristine, recently matured, or restored dunes. San Francisco lessingia is

particularly well-adapted to colonization of disturbed old dunes; see discussion at

Section II.2.a-c and II. 4. for additional information.

Comment: Restoration, as proposed, would require restoration of arid, windy,

unstable dunes that are infeasible in an urban park.

Response: Arid, windy, unstable dunes are not only infeasible in an urban park, they

are contrary to the recommendations of the recovery plan. San Francisco lessingia

does not tolerate significant dune mobility. It is native to gaps in "old growth" dunes,

ancient (even by redwood tree standards) dunes with extensive vegetation and some

soil development. The unstable dunes at Ocean Beach, for example, are not the type

recommended for the Presidio and Fort Funston. The impression that may have been

created by some media accounts that mobile, bare dunes are the goal of the restoration

in the recovery plan was not correct.

Comment: Restoration as recommended would require importing species from other

parts of California, such as Marin County. This is just as arbitrary as removing

non-native species from Monterey (such as Monterey pine).

Response: All of the key species recommended for reintroduction to San Francisco

were historically native associates of San Francisco lessingia, well within natural

geographic ranges, and were only recently driven to localized extinction by urban
Appendix VI - 18

development during the 20th century. The relevant time-frame for ecological

restoration is the recent geological epoch, not earlier geological epochs with radically

different climates, coastline configurations, fauna, and vegetation. Principles for

reintroduction of native species require that seeds and plants are obtained from

appropriate sources intended to minimize the long-distance importing of species or the

introduction of inappropriate species. These principles include obtaining seeds or

individuals from source populations that are as nearby as is feasible, that exhibit

similar genetic and ecological characteristics, and come from similar habitats and

substrates.

Comment: There is no evidence that wind (wind fetch, exposure) is necessary for the

conservation of San Francisco lessingia habitat; therefore, recommendations for the

removal of trees or large restoration sites is unjustified.

Response: The relationship between wind exposure and habitat for dune annuals like

San Francisco lessingia is indirect, based on habitat conditions rather than wind itself. 

Dunes are created by wind, and windflow affects evaporation and transpiration rates

(water loss from plant tissues, physiological aridity). Aridity affects plant growth, and

competition between plant species. Infrequent strong winds can etch out "deflation"

sites in dunes, superficially scouring sand in local patches where vegetative cover is

kept sparse and exposed by episodes of drought. The interaction between wind stress

(that "prunes" vegetation to low-growing forms, and causes physiological aridity) and

wind disturbance (deflation) of otherwise stable dunes is an essential process to the

natural dynamics of dune ecosystems.

Comment: Tree removal in the Presidio will destabilize the dunes and expose

underlying sand to severe wind transport, causing blowing sand hazards comparable to

the Great Highway (Ocean Beach, periodically closed during high winds) and

excessive erosion. This was the original condition of the Presidio before trees were

planted, and the reason why trees were planted.

Response: Most of the Presidio's early historic dunes were "old growth" stable dunes

with scrub and grassland, which were converted to rough pasture during the Mission

era. Extensive, mobile, largely unvegetated dunes occurred landward from Ocean

Beach and Lands End. The historic localities of San Francisco lessingia were from

areas with long history of vegetative stabilization, but were locally disturbed during
Appendix VI - 19

the military period. The recovery plan does not recommend creation of unstable

dunes; it recommends restoration of vegetated dunes with only local gaps, and

establishes criteria to limit sand mobility. San Francisco lessingia is intolerant of

significant sand movement. Dune stability is as readily achieved by native vegetation

as it is by artificial tree plantations; in fact, native vegetation establishes more rapidly

and provides stabilization sooner than tree plantations, which historically were

preceded by other stabilization methods such as brush fencing and planting of

beachgrass and lupine. The technology for stabilizing coastal dunes is old and

well-tested.

Comment: What were the criteria for selecting these species for recovery plans?

Response: We are required by the Endangered Species Act to prepare recovery plans

for all federally listed species unless it would not benefit the species to do so. 

Comment: Several commenters, including the National Park Service, cultural

conservancy groups, and individuals were concerned about a potential conflict

between restoring habitat and meeting the objectives of the National Historic

Preservation Act. One commenter suggested keeping several portions of the historic

forest that the recovery plan recommends for restoration to dune scrub. The

commenter said the stature and spacing of trees could be modified to allow for wind.

Response: We acknowledge that the historic forest is a contributing element of the

Presidio of San Francisco National Historic Landmark District as designated under the

National Historic Preservation Act. Decisions between conflicting resource values are

always difficult to make. By incorporating adaptive management strategies and

engaging in further planning efforts for areas where the historic forest overlaps the

Presidio Recovery Unit, it may be possible to retain the historic character of the forest

while providing optimal recovery of San Francisco lessingia. The intent of the

recovery plan is to provide for the recovery of the San Francisco lessingia, and

therefore, it outlines an aggressive program for that recovery. It is possible that

recovery strategies may result in an adverse effect to the historic forest, but we will

work constructively with the National Park Service and the Presidio Trust to avoid,

minimize, or mitigate any adverse effect to the overall National Historic Landmark

District.
Appendix VI - 20

As part of our efforts to work with the National Park Service and Presidio Trust to also

meet the requirements of the National Historic Preservation Act, we have agreed that

the historic forest area on Rob Hill should be managed as historic forest as a buffer to

the San Francisco lessingia habitat. We have also agreed to management of the

Building 1750 and Bowley Street Interim Management Area as historic forest while

restoration of other areas proceeds. If wind movement is adequately enhanced, it may

be possible to change the recommendation for conversion of this area. We have also

agreed to work together with the National Park Service to prepare a management plan

for the Special Management Zone identified in the Presidio Vegetation Management

Plan. The management plan would evaluate the recovery needs to determine what

changes to the areas of historic forest, if any, are needed for recovery. 

Comment: It may be useful to establish a Memorandum of Understanding between

the National Park Service and the U.S. Fish and Wildlife Service to assist in

cooperation for implementing the recovery plan.

Response: We agree with the National Park Service that a Memorandum of

Understanding to provide for closer cooperation between our two agencies might be

useful in implementing the recovery plan. We recommend including a provision for

the evaluation of endangered species habitat management on National Park Service

lands under this broader view of the Heritage Landscape provisions of the National

Historic Preservation Act.

Comment: San Francisco lessingia should be adequately conserved in a much smaller

area than the recovery plan recommended. The large population sizes of San

Francisco lessingia in the Presidio indicate that small areas should be sufficient to

conserve this species. However, it has been difficult to sustain the intensive

management needed to maintain San Francisco lessingia, so expanding the population

to a larger area would become nearly impossible. 

Response: We believe that the scientific information supports the size of the

recommended conservation areas. The recovery plan reported population data from

the National Park Service which indicated a temporary, unstable, large surge in

population size of San Francisco lessingia in the Presidio soon after major restoration

work (weed removal, tree removal) occurred. This population surge was followed by

a major decline in population size as the wind-sheltered restoration sites rapidly
Appendix VI - 21

developed a relatively dense, continuous vegetation canopy of low shrubs and

perennial herbs, which left little available open sandy sub-habitat for dune annuals like

San Francisco lessingia. This rapid "boom and bust" cycle suggests that sustainable

populations of this species depend not merely on area of habitat, but the essential

environmental processes that influence the quality of habitat. Attempting to conserve

San Francisco lessingia in small areas does not allow for these necessary ecological

processes, increases the probability that these small populations will be subjected to

random deleterious events, and does not fully ameliorate or eliminate the threats that

caused the species to be listed for Federal protection. Population size alone is not the

overriding recovery objective: persistence of the species in its natural role in the

community, maintained by ecosystem processes rather than cultivation, is the ultimate

recovery goal. Small areas under intensive active management could indeed be forced

to produce large numbers of San Francisco lessingia, but this methodology can only

contribute to the species' survival in the short-term; it is not a valid substitute for

recovery.

Comment: The National Park Service commented that the detailed actions and

timelines are overly prescriptive and need to be modified to allow flexibility, while

another commenter requested additional descriptive information about site-specific

management and existing conditions.

Response: We recognize the need to adjust to changing circumstances and new

information. We are fully aware that each step of recovery implementation may not

correspond exactly to what was written in the plan. However, according to our policy,

it is necessary for us to present an integrated, detailed plan and project a period of time

to accomplish the recovery objectives. Projecting a schedule of recovery actions into

the future is actually advantageous for land managers in the sense that it provides a

rational basis for future budget requests. 

The Endangered Species Act of 1973, as amended, section 4, requires description of

actions necessary, including site-specific management actions, to conserve endangered

species. A wealth of site-specific information is available at the Presidio because of a

long history of botanical study, strong natural resource management programs, and

available expertise. These favorable circumstances allowed geographic

recommendations to be formulated. At the same time, a recovery plan cannot serve as

an encyclopedic archive of all information about the species and their sites. A
Appendix VI - 22

recovery plan typically provides enough detail to explain the basic biology of the

species, present a logical explanation for how and why recommendation were

developed, and to provide strategies and criteria for the recovery of listed species. It is

not a regulatory document or a site-specific restoration plan.

Comment: The National Park Service would like us to integrate the environmental

review process required by the National Environmental Policy Act in the recovery

planning process before finalizing the recovery plan to harmonize the recovery actions

with management of the cultural, recreational, viewshed, and other resource values

that the National Park Service is also responsible for.

Response: We recognize implementing a recovery plan may have effects on cultural

and other environmental resources that need to be analyzed under the National

Environmental Policy Act. It is important to remember, however, that a recovery plan

is not a regulatory document. Rather, it delineates actions that we believe are required

to recover and/or protect listed species. For those recommended recovery actions not

categorically excluded, further analysis will be performed under the National

Environmental Policy Act, as specific projects are implemented. We agree that a

cooperative approach between all interested parties is the best approach to managing

lands that are the subject of multiple land use mandates, which are at times conflicting

uses. We also believe that the planning and management process is an ongoing

venture, for which any particular document along the way must seek to improve our

ability to meet the objectives we are charged with. It is our intention to facilitate and

promote a formal coordination and cooperative working group with all interested

parties.

Comment: The National Park Service recommended adding guiding principles to

complement and to clarify our recovery strategies. These guidelines included: (1)

establish conditions that maximize the role of natural processes in sustaining listed

species’ populations and habitat relative to the role of active management; (2) reduce

the cover of invasive non-native plants within the population boundary of listed

species and their habitats; (3) expanding and connecting existing listed species

populations into larger, contiguous suitable habitat should be a higher priority than

establishing smaller, isolated suitable habitats; and (4) habitat restoration should be

implemented so as to restore the listed species’ full communities, including the
Appendix VI - 23

community associates that are “Species of Concern or Regional Conservation

Significance.”

Response: We have implicitly included these guiding principles in developing our

recovery criteria and the actions necessary to achieve recovery. For example, in the

Interim Recovery Criteria for San Francisco lessingia (page vi), we have stated that

the cover of nonnative vegetation in the Lobos Creek, Battery Caulfield, Wherry, Rob

Hill, and Public Health Services Hospital sites should be less than 5 percent, 20

percent, 5 percent, 20 percent, and 20 percent, respectively. Although this interim

recovery criterion is more specific than the stated principle, we believe this criterion

follows the principle of reducing the cover of invasive nonnative plants within the

population boundary of listed species and their habitats. Our recovery criteria are

more specific, because the Endangered Species Act requires us to have objective

measurable criteria which, when met, would result in a determination that the species

can be delisted. However, we recognize that new data may change our appraisal of

what constitute appropriate recovery criteria, and that future modifications of recovery

actions will be informed by research results and management experience that follow

from implementation of recovery actions. 

Comment: The recovery plan must provide an explanation of the reasons that actions

given priority 1 importance are necessary to prevent extinction. This does not seem to

be included in the Stepdown Narrative.

Response: The key section of the recovery plan in this respect is “Section IV.

Recovery.” The recovery strategies presented in this section for San Francisco

lessingia and Raven’s manzanita describe the salient features of the biology and

threats that lead us to recommend the actions found in the narrative outline and

implementation schedule, and outline a general rationale for the priority and order in

which recovery actions should be accomplished. The interim and long-term recovery

criteria also act as benchmarks to measure how the immediate and long-term

conservation needs of the species are being met. We are working on improving the

methods we use in recovery plans to show the connection between threats, actions, and

priorities. To this end, the table in Appendix V links each threat with the specific

recovery actions needed to address it and the recovery criteria that would indicate that

the threat is being adequately ameliorated.
Appendix VI - 24

Comment: Lower the priority of establishing a population at Sutro Heights to one

reflecting its usefulness as a research location over and above its likelihood of serving

as a secure, long-term wild recovery population. Identify the potential resource

impacts of “Rotational Use Areas” recommended for the Fort Funston location.

Response: We recognize that the Sutro Heights site may be more problematic and

may have less likelihood of success than other locations. However, we believe that the

importance of establishing multiple populations is essential for the recovery of San

Francisco lessingia. This is done to address the risks of relying on a single site.

We agree that the different characteristics of this site make it suitable for analysis of

the factors affecting establishment, reproduction, and management. We think this also

applies to the different situation at the Fort Funston location.

Comment: Two separate letters discussed the priority of satellite populations. One

letter suggested the satellite sites should take a high priority for implementation, while

the other suggested a lower priority for these sites.

Response: We have examined and reassessed the priorities of each of the recovery

actions regarding the satellite populations. All of these actions are currently ranked as

priority 1 or 2. Based on our assessment, we feel confident that these actions are

correctly prioritized to meet the conservation needs of the species. 

Comment: One commenter recommended that we obtain independent peer review of

the draft recovery plan by qualified plant population ecologists. Another commenter

felt the draft recovery plan was unscientific and rationalized an agenda to restore

arbitrarily preferred native species habitat.

Response: We followed our policy for peer review (U.S. Fish and Wildlife Service

and National Oceanic and Atmospheric Administration 1994b) and requested peer

review from 16 experts (refer to section A of this appendix). Scientific peer reviewers

provided constructive criticism of the draft recovery plan, recommending areas where

additional explanation would be appropriate. No scientific peer reviewers, with

expertise in the subjects covered, suggested that the draft was unscientific or

rationalized an agenda for unwarranted restoration.
Appendix VI - 25

Comment: The recovery plan does not adequately justify the recovery criteria or

specifications of recovery recommendations; it merely asserts or describes them.

Response: We agree that the draft plan does not offer explanations of how recovery

criteria or recovery strategies were developed. There is some limitation on how far a

planning document can go into these types of explanations. Some explanation is

essential, while other information is implicit in the literature cited by the plan. Our

intention was not to reproduce the articles providing information about recovery needs

and methods, but to apply that information to make scientifically sound

recommendations on the recovery of these species.

Comment: A site-specific plant species list, recognizing the "sub-ecosystem" of each

site, is essential to recovery objectives.

Response: The recovery plan is not an all-inclusive site restoration plan. 

Site-specific restoration plans would be developed by lead agencies during

implementation, in cooperation with us. The plant community, and specific

herb-dominated assemblages in which San Francisco lessingia belongs, was addressed

both in Section II A .3.b. (Plant Associations), III (Species of Concern), and

Appendix I.

Comment: "Delisting" is not an appropriate goal for San Francisco lessingia and

Raven's manzanita.

Response: Delisting is the statutory goal for all federally listed endangered species,

unless it is precluded. For Raven's manzanita we have developed recovery criteria

only for downlisting to threatened status, and not for delisting, because of its

constrained recovery potential. So, for that species, we agree that there is no clear

path that we can anticipate at this time toward independent, secure survival in the wild

without protection of the Endangered Species Act. For San Francisco lessingia, we

believe that delisting is conceptually possible. There appears to be sufficient habitat

available for restoration to support populations that would not need intensive

management if invasive weed species are being managed on a continuing basis in

restored dune scrub habitat.

Comment: Raven's manzanita does not need another recovery plan.
Appendix VI - 26

Response: Our policy is to update older recovery plans when new scientific

information or new circumstances warrant. Much new basic scientific information is

now available about this taxon's genetic relationships, and circumstances in the

Presidio are entirely different than they were when the original recovery plan was

written. The recovery plan was ripe for revision, and the new listing of San Francisco

lessingia provided the occasion for merging the two plans in a single geographic

coverage with related land management.

Comment: Weeding (manual weeding) should be the main task for recovery,

requiring full-time gardeners or field botanists.

Response: Weeding is a useful short-term conservation tactic to conserve existing

populations, but chronic dependence on weeding for a species' survival would indicate

a failure of recovery. The purpose of the Endangered Species Act is to conserve the

ecosystems on which endangered species depend. The recovery plan recommends

restoration of the integrity of the dune vegetation, its physical factors and processes,

and recovery unit area to edge ratios to reduce the re-invasion potential of annual

weeds, especially European grasses, iceplant, and non-native trees. Some sites, such

as Daly City and the Presidio Golf Course roadside site, are recommended for

intensive management rather than ecological restoration because of site size and

irreversible constraints on adjacent land use compatibility.

Comment: There is no mention of San Francisco spineflower (Chorizanthe

cuspidata).

Response: This species was treated under Species of Concern, Section III.

Comment: The U.S. Fish and Wildlife Service needs a consultant familiar with the

Presidio/Daly City sites.

Response: Implementation of recovery plan recommendations will require many

partnerships and contributions of expertise during preparation of site-specific

restoration and management plans. The National Park Service and Presidio Trust have

retained both their own natural resource staff and consultants to assist in natural

resource planning and analysis. 
Appendix VI - 27

Comment: What is the role of botanical garden populations of these species?

Response: Cultivated conservation collections are indeed recommended for these

species (Section IV), but are merely tools to support recovery; cultivation is not a

substitute for recovery of wild populations. See Section II A.5 (Conservation Efforts)

and II. B. 5 (Conservation efforts).

Comment: Is it ethical to reintroduce Franciscan manzanita (Arctostaphylos

franciscana), which is extinct in the wild? Is it wise or ethical to consider

hybridization as a conservation tool?

Response: Consistent with the Center of Plant Conservation's guidelines,

reintroduction will be used to establish populations within historic geographic and

ecological range. Site-specific reintroductions are not prohibited where no historic

documentation occurs. Populations are dynamic, and availability of reintroduction

sites for endangered and rare species is usually limited. Our peer reviewers have

endorsed the experimental approach to reconstituting fertile populations of Raven's

manzanita.

Comment: What is the geologic history of the Daly City site?

Response: The geologic history of the Daly City site is discussed in Section I.B.1.a.

Comment: It is unclear how the seed collection for San Francisco lessingia is to

proceed to maintain a refugial stock at seedbanks.

Response: We have clarified the needs for seed collection in Recovery Action A 5.2.

Comment: There is an indication that the dunes do not have the potential to be

regenerated by physical processes, in the sense that beach sand is not moving into the

relict dune area. It is true in the narrower sense that dune scrub is regenerated

naturally by the cycle of disturbance and plant succession. In order that the recovery

plan more clearly describe what is intended, it would be useful to explain the

definition of regeneration, as it is meant in terms of dune scrub habitat.
Appendix VI - 28

Response: This seems to be a case where there could, indeed, be miscommunication. 

The intended meaning was that of ecological regeneration of a plant community. As

the commenter points out, some readers might construe the meaning as that which

occurs geologically by the movement of material by wind and waves to build dune

landforms.

To clarify our usage in the text of the recovery plan we have added an adverbial or

adjectival descriptor of the context in which we are working to distinguish between

ecological, vegetative, populational, and geological contexts. Ecological regeneration

results in bringing back the aboriginal community of species. Vegetative regeneration

is when an individual plant regrows from lower stems or rootstock. Vegetational and

populational regeneration is when a species regains its original place in the plant mix. 

Geological regeneration is a process where landforms are maintained by balancing the

flows of incoming material with the erosional processes that wear them away, such as

the building of a sandbar in a river, or with the soil-building processes that transform

unconsolidated mineral materials, as when mobile dunes change into relatively stable

dune scrub, and eventually into hills with a mature soil.

Comment: It was noted that we did not give specific information about the conditions

under which San Francisco lessingia can colonize dune scrub. The locations where it

is found growing are secondary blowouts during their restabilization phase, but we do

not describe fully the difference between conditions during an active disturbance event

or period and those during which the San Francisco lessingia is actually growing.

Response: This is in part because of the many different ways that a site can reach a

state that allows San Francisco lessingia to grow. There could be a single disturbance

event during a large storm, or there could be a slower, more gradual type of

disturbance. For example, hikers walking down a path and breaking the surface carpet

of vegetation is a slower, more gradual event. We believe that we adequately

discussed the difference between various conditions under which San Francisco

lessingia grows in the Introduction and Biology of Federally Listed Species sections.

Comment: Watering, fertilizing, and the use of soil amendments would be a problem

at any site that has potential for restoration of dune scrub habitat for recovery of San

Francisco lessingia, not just at the Presidio.
 

Appendix VI - 29

Response: We agree. We will bear this in mind when communicating with any

landowners who have potentially restorable habitat

Comment: The National Park Service and Presidio Trust have many questions and

comments about specific areas, planning objectives, and methods. For instance, the

Presidio Trust and the National Park Service suggest changes to the boundary of the

Presidio Recovery Unit for San Francisco lessingia.

Response: We have made a number of modifications to the recovery plan based on

comments from the National Park Service and Presidio Trust, particularly with respect

to current vegetation management practices and more realistic time and cost estimates

for recovery actions. We in general believe that the boundaries of the Presidio

Recovery Unit are biologically appropriate in the context of the existing and potential

biological and geological environment and the recovery needs of San Francisco

lessingia. However, we have also made some minor changes in the boundaries of the

Presidio Recovery Unit based on site-specific information, comments, and

recommendations provided by the Presidio Trust and National Park Service. We

recognize that the particulars of implementing recovery actions will require further

development of details about specifications of locations, timing, funding, and

management of other resource values, at the Presidio and elsewhere. We also

recognize that close coordination between us, the National Park Service, the Presidio

Trust, and many other interests and partners will be necessary to successfully reach the

recovery goals of this plan. We are committed to working with everyone who will be

involved and to consideration of all interests and concerns. We believe that a shared

commitment by everyone can succeed in promoting workable solutions.

We want to thank everyone who took the time to comment on the draft recovery plan. 

And we hope that you will continue to be involved to make sure that our efforts to

carry out the plan will be well informed and address problems in a collaborative,

creative, and positive spirit.