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 Investigation of Wind Pollination in the Humboldt BayWallflower (
 Erysimum menziesii 
ssp.
eurekense
) at theLanphere-Christensen Dunes
Joshua Der, Sarah Jordan, Veronica VegaSenior Thesis in BiologyHumboldt State University, Arcata, CA 95521May 2003Final report submitted to Humboldt Bay National Wildlife Refuge.
Abstract
We examined wind pollination as a potential contributor to the pollination success of the Humboldt Bay wallflower. We selected 15 plants each on dune ridges and in dune hollows to test the effects of different wind regimes. Plants were emasculated and caged to excludeinsect pollinators and ensure that wind was the only mechanism availablefor pollination. Stigmas were collected five days after emasculation and pollen loads were counted. We found no significant differences in thenumber of 
 Erysimum
or heterospecific pollen grains, or the ratio of eachon the stigmas between ridges and hollows. There may not have beenenough of a difference between treatments to affect pollen deposition. Wefound more pollen in both treatments than can be attributed tocontamination from emasculation. This suggests that wind may play arole in the pollination system of 
 Erysimum
.
 
Wind Pollination in the Humboldt Bay Wallflower2
Introduction
Plants use a variety of mechanisms to achieve pollination. These pollinationsystems are variable, cryptic, and often poorly understood in most plants. Plants oftenutilize more than one pollination system in order to ensure pollination success andincrease fecundity (Whitehead, 1968; Goodwillie 1999). Ambophily is a pollinationsystem that uses both insects and wind as vectors of pollen transfer (Culley et al., 2002).Ambophily has evolved in several unrelated families of plants including the Asteraceae,Brassicaceae, Ericaceae, Salixaceae and others (Culley et al., 2002 and referencestherein). Recent research suggests that ambophily is a transitional phase betweenexclusively animal and wind pollinated systems (Culley et al., 2002). Ambophily may beselected for by low insect visitation, high winds, and semelparity (Goodwillie, 1999).
Wind pollination is generally considered to be a primitive pollination system, andthe diversification of angiosperms has been associated with the evolution of insects aspollinators. However, m
orphological adaptations of flowers in some animal pollinated plants may promote wind pollination. Flattened or reflexed corolla lobes and exerted or elongating stigmas are adaptations which increase the likelihood of wind pollination(Goodwillie, 1999). The contribution of wind pollination to reproductive fitness varieswidely between different plant groups. The proportion of the total pollen transferred bywind varies from 8% to as much as 93% (Culley et al., 2002).The Humboldt Bay wallflower (
 Erysimum menziesii
ssp.
eurekense
, Brassicaceae)is a small Endangered herb endemic to the windy coastal foredunes near Humboldt Bay,California. This habitat faces increasing pressure from off road vehicle use and invasive plants.
 Erysimum
 produces a small inflorescence as early as mid-February, when pollinator abundance is low. The flowers are small and have many characteristicscommonly associated with bee-pollinated plants: showy yellow flowers, abundant pollen,nectar rewards, and petals that provide a landing platform for bees.
 Erysimum
also hasflattened petals and an elongated style exposing the stigma, increasing the opportunity for wind pollination. Although insect visitation is limited, fruit and seed set are nearlycomplete (M. Mesler and J. Sawyer, unpublished). These factors suggest that
 Erysimum
may utilize wind as a vector of pollination in addition to insect pollination.
 
Wind Pollination in the Humboldt Bay Wallflower3
Understanding the reproductive biology of this Endangered plant, allows us todevelop effective management strategies to conserve the species and preserve theintegrity of its rare community. Since there has been little research on ambophily to date,it is important to examine this as a potential mechanism of pollination in plantstraditionally thought to be exclusively insect pollinated.The purpose of this study was to determine if 
 Erysimum
utilizes wind pollinationas part of its pollination system. We expected to find a higher rate of pollination inexposed areas than in sheltered areas when we emasculated plants and excluded insect pollinators.
Methods
Study Species and Site
 Erysimum
 
menziesii
ssp.
eurekense
is a Federally and State Endangered species,endemic to the Humboldt Bay area. It is found only on the Lanphere and the Samoadunes. It typically flowers between mid February and mid June, with peak flowering between March and April. Insect and self-pollination have both been observed in
 Erysimum
. An average of 29 flowers are produced by an inflorescence over its lifetimeand these flowers open acropetally at a rate of 2.2 flowers per day (M. Mesler and J.Sawyer, unpublished). Plants usually display inflorescences 10-25 cm tall, and larger  plants sometimes branch. Plants are monocarpic and inflorescences bolt from a basalwhorl of leaves after 1-2 years of vegetative growth.
 Erysimum
’s most abundant insect pollinator is the solitary bee,
 Habropodamisirabilis
(M. Mesler and J. Sawyer, unpublished). Other important pollinators include
 Bombus melanopygus, B. mixtus, B. vosnesenskii
and
 Lasioglossum pavanotum
.Occasionally ants and Elaterid beetles will visit the flowers.The Lanphere Dunes Preserve is a unit of the Humboldt Bay National WildlifeRefuge located west of Arcata in Northern California and is characterized by two generalhabitat types: foredunes and dune forest. The foredunes are characterized by a series of dune ridges and hollows.
 Erysimum
occurs predominantly in exposed sandy areas on theforedunes in the northern section of the preserve and appears to have a clumpeddistribution (personal observation). The preserve is an important site that protects these
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