(DRAFT) - Taxonomy
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
TAXONOMY
NAME - EVENING-PRIMROSE, ANTIOCH DUNES
OTHER COMMON NAMES - EVENING-PRIMROSE and ANTIOCH DUNES
ELEMENT CODE -
CATEGORY - Angiosperm
PHYLUM AND SUBPHYLUM - MAGNOLIOPHYTA,
CLASS AND SUBCLASS - MAGNOLIOPSIDA,
ORDER AND SUBORDER - MYRTALES,
FAMILY AND SUBFAMILY - ONAGRACEAE,
GENUS AND SUBGENUS - OENOTHERA,
SPECIES AND SSP - DELTOIDES, SSP.
SCIENTIFIC NAME - OENOTHERA DELTOIDES SSP.
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Antioch Dunes Evening-primrose
Oenothera deltoides ssp. howellii (Munz) W. Klein
KINGDOM: Plant GROUP: Angiosperm
DIVISION: Magnoliophyta CLASS: Magnoliopsida
ORDER: Myrtales FAMILY: Onagraceae
The Antioch Dunes evening-primrose is a perennial species
characterized by few to several decumbent branches reaching 0.5 - 3 dm
high. The plant generally is glabrous below and spreading pubescent
in upper parts. The leaves are pinnatifid, 3-12 cm long and ash gray
in color. The lower leaves form a loose rosette while cauline leaves
are gradually reduced in size. The buds are long-pointed with few to
many wavy hairs. Flowers are solitary in upper axils with the petals
white and fading pink. Members in the genus Oenothera get their
common name, evening-primrose, because the flowers are vespertine,
that is they open in the late afternoon and bloom through the evening.
The seed capsules are woody with an exfoliating epidermis. The seeds
are narrowly obovoid, light brown in color usually with dark spots.
This subspecies is found naturally only in sand dune areas of eastern
Contra Costa County, California (01).
Type specimens are found at the Herbarium, University of
Taxonomy - 1 (DRAFT) - Taxonomy
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
California, Berkeley (01).
The following is taken from the species Recovery Plan:
"The Antioch Dunes evening-primrose is a member of the
evening-primrose family (Onagraceae) and the subgenus Anogra.
Although there is a strong consensus today on the taxonomic
status of the plant ..., this was not always the case. In
1925 Jepson described O. trichocalyx var. cognata, from
Corral Hollow in Alameda County. Munz in 1931 placed a
collection of Oenothera from the Antioch Dunes in Contra
Costa County with O. deltoides var. cognata. This was a new
varietal combination employing Jepson's earlier varietal
epithet. Jepson (1936) later concurred that the material
from Antioch was the same as that from Corral Hollow, but he
did not accept Munz' specific epithet for the Antioch Dunes
evening-primrose.
"Alice Eastwood and John Thomas Howell collected the Oenothera
from Antioch in 1936 and later Howell brought it to the
attention of Munz. Munz concluded from observations of
cultivated and field-collected specimens that the Antioch
Dunes evening-primrose was indeed a distinct entity. He
named the plant O. deltoides var. howellii in honor of Howell.
Munz employed the pinnatifid leaves, long-pointed buds, and
perennial habit to distinguish this evening-primrose from
O. deltoides var. cognata. Later Klein (1962), after a
biosystematic investigation of several entities of the
subgenus Anogra, accorded the Antioch Dunes evening-primrose
along with other taxa subspecific rank. Ultimately Munz
(1965), in his monograph of the Onagraceae in the North
American Flora Series, concurred with Klein's subspecific
treatment of the plant. Furthermore, breeding studies by
Klein (1970) substantiated that the Antioch Dunes
evening-primrose has a strong affinity for the O. deltoides
complex. Consequently O. deltoides subsp. howellii remains
the most appropriate scientific epithet at this time for the
Antioch Dunes evening-primrose .... "(04).
Type specimen - 19 April 1936, Eastwood & Howell 2147:RSA;
isotypes: CAS,POM. Other collections include: Antioch, C.F. Baker
2873 (POM), P.A. Munz 12201 (RAS), V Rattan in 1879 (DS) and K.
Brandagee in 1892 (DS,UC) (07).
Taxonomy - 2 (DRAFT) - Status
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
STATUS
Coded Status
E: Federal Endangered
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The Antioch Dunes evening-primrose (Oenothera deltoides ssp.
howellii) has been designated an Endangered species pursuant to the
Endangered Species Act of 1973 (50 CFR 17.12; P.L. 93-205, 87 Stat.
884; 16 U.S.C. 1531-1540), as amended. The species has this status
wherever found including the State of California. Critical Habitat
has been designated in Contra Costa County, CA (50 CFR 17.96(a)) and
is defined as an area of land, water and airspace in T2N R2E, SW 1/4
of Sec.17 and the E 2/3 of the S 1/2, Sec.18.
Removal and reduction to possession of any Federally listed plant
from an area under Federal jurisdiction is unlawful (50 CFR 17.61 and
17.71).
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: California
DESIGNATED STATUS: Endangered
ADMINISTRATIVE AGENCY: California Department of Fish and Game
STATE STATUTES: California Endangered Species Act; Fish and Game
Code, Chapter 1.5, Article 1, Section 2050.
The California Environmental Quality Act (PRC 2100 et seq.)
recognizes Federally listed Threatened and Endangered species as among
those species requiring that environmental impact assessments be made
for actions that may detrimentally affect them. See: Guidelines for
Implementation of the California Environmental Quality Act, California
Administrative Code, Chapter 3, Section 15380.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
None.
Status - 1 (DRAFT) - Status
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
ECONOMIC STATUSES:
In 1979 the Antioch Dunes evening-primrose was featured on a U.S.
commemorative four-stamp series called "Endangered Flora".
76/06/16:41 FR 25423/25472 - Proposed rule, listing as Endangered
77/02/08:42 FR 07972/07975 - Proposed rule to designate CH
78/04/26:43 FR 17910/17916 - Listed as Endangered
78/08/31:43 FR 39042/39044 - Determination of Critical Habitat
83/12/08:48 FR 55100/55102 - Notice of five year review
85/07/22:50 FR 29900/29901 - Five year review completion
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
LAND USE -
Industrial
Transportation, communications, and Util
Orchards, Groves, Vineyards, Nurseries,
Sandy Areas other than Beaches
Strip Mines, Quarries, and Gravel Pits
Transitional Areas
COMMENTS ON HABITAT ASSOCIATIONS -
The Antioch sand dunes comprised a unique ecological area from
several standpoints, although they are now largely decimated by
industrialization. Biologically Antioch Dunes was a kind of "island"
that contained the northern extension of many plants and animals of
desert affinities; a biogeographic element that probably extended
along the western margin of the Central Valley in prehistoric times.
Later natural processes reduced this biota to a few small islands of
sand dunes, the largest of which was at Antioch. Although the dunes
resemble coastal dunes, such as at Pt. Reyes, the plants and animals
were quite different, being desert species. The long isolation from
relatives in the Mojave resulted in considerable local
differentiation, or endemism, with the occurrence of species or races
peculiar to the one area (08). The dunes once encompassed about 500
acres with elevations to ca. 100 ft above sea level but were reduced
to about 60 acres with elevations of ca. 30 ft above sea level.
The Antioch Dunes evening-primrose is associated with riverine
sand dunes located on Brannon Island State Park, Sacramento River and
along the San Joaquin River. Portions of the Antioch Dunes population
are located on the Pacific Gas & Electric (PG & E) right-of-way
adjacent to the Antioch Dunes National Wildlife Refuge and is managed
by the Refuge. With establishment of the Antioch Dunes NWR, 55 acres
of mined and unmined sand dune areas were set aside to protect this
species. Average high temperatures in the area range from above 90
degrees F in July, August and September to the low-mid 50's in
December and January. Average low temperatures range from the low
60's (degrees F) in July, August and September to the mid-upper 30's
in December and January. In 1984 rainfall was less than nine inches
for the calendar year with the greatest amount (3.80 inches) falling
in November. Typically, the months of May through September are dry
with little or no precipitation, while the "rainy season" runs from
late October through March.
The Antioch Dunes evening-primrose grows strictly in sandy soil
(Oakley or Delhi sands) with various amounts of silt and/or clay mixed
in. The soil types are classified as sand to sandy loam having a pH
averaging 6.3 (range 5.6-7.0). Currently the evening-primrose is
found where ground cover ranges from less than 5% to 100%. Seedling
establishment appears best where ground cover is minimal. A
particular slope or aspect does not seem to be a specific
environmental paramenter required by this species.
Currently, dune habitat is adjacent to a sewage treatment
facility, old and ongoing sand quarrying areas, an old vineyard, the
Domtar gypsum plant, the Atchison/Topeka and Santa Fe Railroad tracks,
the Fulton shipyard road, and traversed by PG & E power links.
Habitat Associations - 1 (DRAFT) - Food Habits
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
FOOD HABITS
TROPHIC LEVEL -
AUTOTROPH
Food Habits - 1 (DRAFT) - Environment Associations
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Coastal Features: Vegetated offshore islands
G
G
G
G
Environment Associations - 1 (DRAFT) - Life History
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
LIFE HISTORY
HABIT:
The Antioch Dunes evening-primrose is an herb (01).
LIFE CYCLE:
Perennial (04). One of the characters employed to separate
Oenothera deltoides ssp. howellii from O. d. ssp. cognata is its
genuine perennial habit. Differing opinions exist as to the longevity
of the Antioch Dunes evening-primrose. Munz in his descriptive
article in 1949 stated that it was "apparently [a] short-lived
perennial." Roof described the Oenothera as a biennial in
cultivation. Seedlings after blooming their first spring, bloomed
"profusely" the second year. Nonetheless, Roof noted that a few
individuals flowered in later years, but these were "sickly and
worthless as flowering specimens".
Klein stated the perennial condition of the Antioch Dunes
evening-primrose was of adaptive significance. He postulated that the
perennial habit was in response to "more mesic conditions." He
further noted that the manner in which Oenothera deltoides ssp.
howellii perennated was quite distinctive from other members of the
subgenus Anogra (04).
TYPE OF REPRODUCTION:
Sexual (01).
REPRODUCTIVE PHENOLOGY:
Little is known about the reproductive phenology of the Antioch
Dunes evening-primrose. Germination appears to occur from November to
February. Budding begins in March and continues through May (peak).
Flowering runs from April through June (peak). One individual plant
was observed blooming to December. Fruiting occurs from May through
July (peak). Fruit is dispersed beginning in June and runs through
September (peak) (05). The evening-primrose is vespertine; the
flowers open in early evening and close by mid-morning (04).
SEX OR SPORE STATUS:
Monoclinous (05).
POLLINATION, SPORE AND SEED DISSEMINATION:
This Oenothera is self-incompatible and thus requires
cross-pollination for sound seed. Numerous smooth buff- to
black-colored seeds are produced in the capsule (04). Pollination is
facilitated by insects, thought to be beetles and bees (Sphecodogastra
aberrans). Hawk-moths were not known on the dunes until 1983 but have
been reported as pollinators of other Oenothera. Their role in
pollination has not been documented. A number of different insect
species have been observed visiting the plant and leaving covered with
pollen. Seed is disseminated by wind (05). Much information still
needs to be gathered on dispersal mechanisms, patterns and distances.
SEED BIOLOGY:
No scientific studies have been carried out on this subject.
Life History - 1 (DRAFT) - Life History
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
POPULATION BIOLOGY:
No scientific studies have been carried out on this subject for
this subspecies.
In cultivation, seedlings of the evening-primrose develop
vigorously the first spring to a foot or more in diameter. Although
the plant may bloom the first year, it blooms more profusely the
second year. The plants may grow to approximately three feet in
diameter by the second year and can reach nearly 3 feet in height.
After producing an abundance of seed capsules, most of the plants
grown in cultivation die off during their second winter. However, the
supspecies is best regarded as a perennial, albeit probably
short-lived.
The Oenothera seems to prefer growing in nearly pure sand and
should be regarded as a psammophyte. Observations of cultivated
plants suggest that seedlings of the Antioch Dunes evening-primrose
will not reach maturity where adult plants have recently matured and
died. Fresh sand may be necessary for seedling re-establishment in
such areas. It was speculated that the evening-primrose exhausted the
nutrients within the soil where the preceding generation matured.
However, other factors of cultivation, like watering regime, planting
depth, allelopathy, and disease, might have led or contributed to the
death of succeeding generations of cultivated Oenothera seedlings.
Despite abundant seed production, few seedlings of Oenothera
avita ssp. eurekensis were seen on the Eureka Dunes. Germination of
psammophytes may be dependent on the optimal coincidence of at least
three factors: burial, moisture, and critical temperature regime.
Antioch Dunes evening-primrose has been observed to produce numerous
seedlings in recently disced areas. This might indicate that the
seeds respond favorably to disturbance or burial and not necessarily
the deposition of nutrient-laden fresh sand as postulated.
Regardless, the data are weak on the population biology of Oenothera
deltoides ssp. howellii and further studies are needed for proper
management of the dunes and the subspecies (04).
ECOLOGICAL/EDAPHIC FACTORS:
Information about the species particular requirements are not
known at this time. According to Atwater (02) the dunes were of
aeolian origin from glacially derived sediments laid down during the
late Pleistocene in the Sierra Nevada. Carpenter and Cosby (03)
mapped this area as Oakley sands. Soil types at the refuge have been
classified as ranging from sand to sandy-loam; pH averages 6.4 (range
5.6-7.0). Average high temperatures in the area range from above 90
degrees F in July, August and September to the mid 30's F in December
and January. Typically, the months of May through September are dry,
while the "rainy season" runs from late October through March. In
1984 rainfall was less than nine inches for the calendar year with the
greatest amount (3.80 inches) falling in November.
TROPHIC STATUS:
Phototrophic (01).
CHARACTERISTIC DOMINANCE:
Life History - 2 (DRAFT) - Life History
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
Component (04).
COMMUNITY ECOLOGY
The Antioch Dunes evening-primrose generally is associated with
early successional stages leading up to an oak woodland climax
habitat. The associated plant community includes some of the
following native species: Clarkia unguiculata, Eriogonum nudum,
Eschscholzia californica, Gutierrezia divergens, Lotus scoparius,
Lupinus albifrons, L. bicolor, and Senecio douglasii. Since
acquisition of the land for a refuge, the area has had disturbances
associated with fires, fire control, habitat management and
unauthorized entry by people and their bicycles/motorcycles. Their
disturbances obviously kill any plants growing in the area at the time
of the disturbance, however, for the most part the endangered species
do repopulate the area quickly if the disturbance is not too frequent
or repeated. Some areas of the refuge are actively managed to be kept
in early successional stages for the benefit of the Endangered species
(04,05).
SPECIES INTERRELATIONSHIPS:
Infestations of beetles (possibly Chrysomelid), which attach to
leaves and flower buds, have been noted to reduce reproductive output.
Beetles feed on petals, pollen, and seed capsules. This may need
further investigation as the beetles may be a serious pest.
The evening-primrose is probably pollinated by bees
(Sphecodogastra aberrans), beetles, and/or hawkmoths.
OTHER LIFE HISTORY DESCRIPTORS:
Oenothera deltoides ssp. howellii has extensive fleshy roots
which penetrate to considerable depths into the sand and buds located
near the crown in contrast to the creeping horizontal stems of
O. avita ssp. avita and O. a. ssp. eurekensis (04).
Life History - 3 (DRAFT) - Management Practices
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Suppressing wildfire
Beneficial Controlling/Restricting Off-Road Vehicles
Beneficial Restricting/regulating human disturbance of populations
Beneficial Controlling/Restricting Mining
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Land Acquisition
Beneficial Reforestation
Beneficial Controlling/Removing Nonnative Vegetation
Beneficial Stocking captive-reared wild-strain animals
Beneficial Transplanting Wild Eggs/Wild Seeds
Adverse Off Road Vehicles
Existing Off Road Vehicles
Adverse Hiking/Camping
Existing Hiking/Camping
Adverse Predation
Existing Predation
Adverse Surface Mines
Existing Surface Mines
Adverse Rural Residential/Industrial Areas
Existing Rural Residential/Industrial Areas
Adverse Transmission Lines/Towers
Existing Transmission Lines/Towers
Adverse Soil compaction by heavy equipment in mine areas
Existing Soil compaction by heavy equipment in mine areas
Adverse Competition
Existing Competition
Adverse Exotic/Feral/Introducted Species
Existing Exotic/Feral/Introducted Species
Adverse
Existing
Adverse Vegetation Composition Changes
Existing Vegetation Composition Changes
Adverse Suppressing wildfire
Existing Suppressing wildfire
COMMENTS ON MANAGEMENT PRACTICES -
The Antioch Dunes evening-primrose is endemic to the Antioch
Dunes, which once extended for about 3 miles parallel to the
Sacramento - San Joaquin Rivers. Remnants of the dunes and the
evening-primrose itself now are limited to an area of no more than 60
acres.
Loss of habitat, primarily through sand mining, was a key factor
contributing to its decline. Industrial development, periodic discing
for fire control, hiking, and off-road vehicle use also have had an
adverse affect on the habitat. Of these impacts on habitat only
fuelbreak discing remains a threat both directly as well as indirectly
by increasing the weed species. Predation by beetles (possibly
Chrysomelids) upon petals, pollen, and seed capsules has been reported
Management Practices - 1 (DRAFT) - Management Practices
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
as a serious threat to the species (04). An increase in the number of
weeds, by invasion of exotic grasses, also has stabilized some dune
slopes, and competitively reduced the amount of area available to the
evening-primrose for colonization. Pacific Gas & Electric (PG & E)
maintains transmission lines on two parcels that it owns. Routine
maintenance procedures generally cause little damage to the habitat,
but downed power lines and their repair have caused several fires and
the destruction of the native vegetation by heavy equipment (09,10).
A vineyard, established over 100 years ago, also contributed to
habitat loss.
With the establishment of the Antioch Dunes National Wildlife
Refuge, 55 acres of mined and unmined sand dune areas were set
aside to protect this species, two other listed Endangered species, a
number of species that are candidates for listing under the Federal
Endangered Species Act, and the State-listed California legless
lizard.
Not all the Antioch Dunes evening-primrose plants are located on
refuge lands. One large stand of evening-primrose is found on a
Pacific Gas & Electric (PG & E) right-of-way parcel adjacent to the
refuge. However, this and other PG & E right-of-way parcels are being
managed by the U.S. Fish and Wildlife Service under a cooperative
agreement. An introduced population of Antioch Dune evening-primrose
is found in Brannon Island State Park, approximately ten miles
upstream on the Sacramento River (04,05).
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1984. Revised Recovery Plan for
Three Endangered Species Endemic to the Antioch Dunes, California.
Approved: March 21, 1980 and Revised: April 25, 1984. U.S. Fish and
Wildlife, Portland, OR. 66 pp.
The Recovery plan for the Antioch Dunes ecosystem recommends:
1. Protect the Antioch Dunes ecosystem by:
A. Developing protective alternatives and actions in a
management plan for refuge land and implementing the
management plan. The management plan should include
maintenance of habitats (protection from development such as
mining), control or management of public use (off-road
vehicles and human access to important habitats), and
measures to control or manage fuelbreaks and other fire
control activities;
B. Developing protective alternatives and actions with private
land owners by developing MOU with landowner of essential
habitat; and
C. Identifying other essential habitat for protection.
2. Restoring the Antioch Dunes ecosystem, increasing populations of
plants and improving habitat by:
A. Conducting annual census of population and monitoring
habitat;
B. Propagating plants and outplant (if necessary); and
Management Practices - 2 (DRAFT) - Management Practices
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
C. Developing and implementing a habitat restoration plan based
on complete life-history studies (repoduction history and
autecological studies). (Although not specified in the
recovery plan, transplanting wild seed to restored dune
habitat is recommended.)
3. Rebuilding the natural dune substrate and topography to the
degree feasible using sandy dredge spoils. Reduction of impacts
on candidate species and other listed species which may be
affected by spoils must be considered. Monitoring of site
renovation will be necessary.
4. Reducing or eliminating exotic vegetation.
5. Initiating an information and education program such as
A. Erecting interpretive signs;
B. Printing and distributing leaflets explaining the need for
the refuge and restoration of the dune ecosystem; and
C. Developing an environmental education program.
Essential habitat has been protected through acquisition of key
lands into the National Wildlife Refuge System. The dune's ecosystem
is being restored to improve habitat and increase populations of the
Antioch Dunes evening-primrose, and wild seed has been transplanted to
some restoed sites.
Management Practices - 3 (DRAFT) - References
Species EVENING-PRIMROSE, ANTIOCH DUNES
Species Id ESIS701019
Date 13 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Munz, Philip A. 1968. A California flora and supplement.
University of Calif. Press, Berkeley. 1925 pp.
02 Atwater, Brian F. 1982. Geologic factors in the prehistoric
establishment of organisms endemic to aeolian dunes near Antioch,
CA. Memo prepared for USFWS. 6 pp.
03 Carpenter, E.J. and S.W. Cosby. 1939. Soil Survey, Contra Costa
County, California. USDA, Bureau of Chemistry and Soils.
04 U.S. Fish and Wildlife Service. 1984. Revised Recovery Plan for
Three Endangered Species Endemic to Antioch Dunes, California.
Approved: March 21, 1980 and Revised: April 25, 1984. U.S. Fish
and Wildlife Service, Portland, OR. 66 pp.
05 Robinson, Lee. 1985. Personal observation. 19046 Mayberry Drive,
Castro Valley, CA 94546.
06 Nichaus, Theodore. 1977. Rare plant status report. Calif. Nat.
Plant Soc. Unpulished report on file at: U.S. Fish and Wildlife
Service, Office of Endangered Species, Washington, D.C.
07 Munz, P.A. 1949. California misscellany - I. Aliso 2:81-83.
08 Powell, J.A. and R.A. Arnold. 1977. Status of certain insect and
plant survivors of the Antioch sand dunes. Unpublished report on
file at: U.S. Fish and Wildlife Service, Office of Endangered
Species, Washington, D.C. 3 pp.
09 Arnold, R.A. 1984. Monitoring of the endangered Lange's metalmark
butterfly in 1983 at the Antioch Dunes. Annual Rept. to San
Francisco Bay NWR.
10 Arnold, R.A. 1985. Ecological studies of the endangered Lange's
metalmark butterfly at the Antioch Dunes. Annual Rept. to San
Francisco Bay NWR.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 U.S. Fish and Wildlife Service. 1984. Revised Recovery Plan for
Three Endangered Species Endemic to Antioch Dunes, California.
Approved: March 21, 1980 and Revised: April 25, 1984. U.S. Fish
and Wildlife Service, Portland, OR. 66 pp.
References - 1