Broad
spectrum herbicides such as hexazinone are sometimes applied after
forest fires to increase growth and survival of conifer seedlings
by reducing competing vegetation. This practice removes not only
undesirable alien plant competitors, but also native species.
This effect may be of special concern for rare native species
such as Iliamna bakeri (Malvaceae), a sensitive plant of
northern California. This plant is adapted to follow fire in early
successional stages, and therefore, may be especially vulnerable
to post-fire herbicide application. Currently, USDA Forest Service
land managers have no basis for evaluating this risk because little
is known of the biology of I. bakeri. I and my collaborators
are currently examining the effects of hexazinone application
on the demography, reproduction, and population genetic structure
of I. bakeri. Based on the findings, site-specific recommendations
will be developed regarding hexazinone application, to achieve
necessary levels of conifer growth while supporting the reproduction
of this sensitive plant species. Additionally, we will be examining
the pollination relationships of I. bakeri. Preliminary
observations (S. Smith, USDA) indicate that a specialist bee,
Diadasia nitidifrons, may be the primary pollinator of
I. bakeri. Specialized ecological interactions such as
this must be understood, and management decisions must address
the needs of the pollinator as well as those of the rare plant
species. The relationship of I. bakeri and D. nitidifrons
also presents an optimal opportunity to examine the efficacy of
specialist bees as pollinators of their host plants. Photo credit
Charles Webber, California Academy of Sciences.
Collaborators:
Vincent J. Tepedino, USDA/ARS Bee Biology and Systematics Laboratory, Logan UT.
Sheri L. Smith, USDA Forest Service, Susanville CA.
Spiranthes
diluvialis is a terrestrial riparian orchid that occurs in
widely scattered populations in the western mountains of the U.S.
The species was initially believed to be quite rare and declining
in abundance, and in 1992 it was listed as threatened under the
U.S. Endangered Species Act. Since that time, an heroic surveying
effort by various land management agencies and private interests
has shown it to be more widespread than initially supposed. Nevertheless,
this species' continued survival remains threatened by human modification
of its riparian habitiat (dam building, flood control). The orchid
is an early to intermediate successional species and depends largely
on natural flood disturbance to maintain suitable habitat. I have
been studying the reproductive ecology of S. diluvialis
since 1992, with field studies still ongoing. My research has
focused on:
1) Breeding system and pollination. By experimental hand-pollinations in the field, I determined that, although S. diluvialis is physiologically capable of self-pollination, it is predominantly outcrossing due to the acropetal development of the inflorescence, and the functional protandry of the flowers. Long-tongued bees of the genera Bombus and Anthophora (the primary pollinators) tend to forage from the bottom up on vertical spike inflorescences, thus the bees most often transfer pollinia from the top, younger male-phase flowers of one plant to the bottom, older,female-phase flowers of the next plant visited. However, there is considerable overlap in the sexual phases of flowers, and the specific characteristics of this overlap make self-pollinations especially likely under conditions of low pollinator visitation. Thus, there is a threshold pollinator abundance below which the outcrossing mechanism of S. diluvialis fails to perform. Land managers must strive to protect this rare plants pollinators, and maintain appropriate habitat for the bees.
2) Long-term studies of the correlates of male and female reproductive success. I and my collaborators have measured spatial and temporal variations in reproductive success over a period of 9 years. Because S. diluvialis pollen occurs in a single discrete packet (a pollinarium), the presence or absence of pollinaria serves as an estimate of male reproductive success. Thus we have been able to estimate both male reproductive success (pollinarium removal) and female reproductive success (fruit and seed set).
3) Experimental transplantation of pollinators. Our long-term reproductive studies showed that a disjuct population of S. diluvialis in Daggett co. UT, consistently suffered extremely low rates of bee visitation, pollinaria removal, and fruit set; these data strongly suggest pollinator-limited reproduction. In an attempt to augment pollination levels, my collaborators Vince Tepedino and Kim Pierson and I, working with federal land managers, have trapped and transplanted a native, solitary, twig-nesting bee pollinator (Anthophora terminalis) from sites in Utah co. to the Daggett co. sites. It is too early to know if these actions will significantly increase S. diluvialis reproduction in the long run, but preliminary data indicate a slight reproductive increase in the year following the release of transplanted bees. This novel conservation approach represents the first time native solitary bees have been transplanted in the management of a rare plant species.
Collaborators:
Publications and Research Reports:
As
a side project to my M.S. research, I examined the pollination,
breeding system, and population structure of Jones cycladenia,
Cycladenia humilis var. jonesii, a federally-listed
threatened, edaphically-restricted species occurring in just a
few locations in Utah and Arizona. This taxon exhibits extremely
low fruit and seed set even with hand-pollination, and no seedling
recruitment has been observed. However, the taxon does reproduce
clonally. Population genetic structure inferred from isozymes
revealed unusually high levels of genetic variation and little
evidence of inbreeding, and confirmed that populations consist
of numerous interdigitated clones. I hypothesize that this taxon
may be a paleoendemic surviving in the absence of its original
pollinator through clonal growth. The high genetic variability
may suggest that high genetic load has accumulated in very old
clones, which in turn may also play a part in its current low
levels of sexual reproduction. In an effort to further understand
the restricted distribution of this species, I collaborated with
soil scientist Janis Boettinger (USU) to study the edaphic relationships
of C. h. var. jonesii. This species occurs only
in certain members of the Chinle, Cutler, and Summerville formations
in southern Utah and Northern Arizona. Because these soils are
often gypsum rich, previous reports had classified Jones' cycladenia
as an obligate gypsophile. However, we concluded that the physical,
rather than chemical, characteristics of the plant's soil substrate
are associated with its restriction to only certain geological
members. Jones cycladenia sometimes occurred in very low-gypsum
soils, but all soils in which it occurred shared numerous common
physical attributes (i.e. shallow soils derived from blocky-fracturing
mudstones with high % of rock fragments), and these characteristics
were lacking in adjacent soils where Jones cycladenia did not
occur. These studies have provided important information for land
managers: sexual reproduction is rare and thus does not represent
a means to augment population size or number. Additionally, the
edaphic restrictions limit potential sites for new populations,
so existing populations of this taxon must be protected. Photo:
Jones cycladenia flower bud marked for monitoring, photo credit
Sedonia Sipes.
Collaborators:
Publications and Research Reports:
Pendland's beardtongue
(Penstemon penlandii) is a very rare species in the snapdragon
family (Scrophulariaceae) that occurs in only one county in central
Colorado. The species is listed as endangered under the U.S. Endangered
Species Act. Its continued existence depends critically on being
able to produce sufficient fruit and seed so that new individuals
will be available in the future to replace adults in current populations
when they die. We studied the reproductive biology and pollinators
of Penland's beardtongue to determine if pollinators are required
for fruit set, and to learn what those pollinators might be. We
learned that plants reproduce only minimally in the absence of
pollinators, and that the important pollinators are numerous species
of megachilid bees in the genus Osmia, and bumblebees (Bombus
spp.). Interestingly, the pollinator guild included over a dozen
species of Osmia, which occur sympatrically within P. penlandii's
habitat. However, in any given year, only a few Osmia species
might be abundant. If Penland's beardtongue is to be preserved
then its pollinators must also be encouraged to thrive. Because
of the complexity of the pollination system in which it is embedded,
this plant is best preserved by applying principles of ecosystem
conservation. Photo copyright B. Jennings.
Collaborators:
Publications:
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