Phacelia argillacea - Atwood
Clay Phacelia
Taxonomic Status: Accepted
Related ITIS Name(s): Phacelia argillacea Atwood (TSN 31455)
Unique Identifier: ELEMENT_GLOBAL.2.133894
Element Code: PDHYD0C080
Informal Taxonomy: Plants, Vascular - Flowering Plants - Waterleaf Family
 
Kingdom Phylum Class Order Family Genus
Plantae Anthophyta Dicotyledoneae Solanales Hydrophyllaceae Phacelia
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Phacelia argillacea
Conservation Status
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NatureServe Status

Global Status: G1
Global Status Last Reviewed: 15Nov2012
Global Status Last Changed: 15Jul1983
Ranking Methodology Used: Ranked by inspection
Rounded Global Status: G1 - Critically Imperiled
Reasons: A narrow endemic of Utah County, Utah. There are only two known populations. Since this species is a winter annual and is extremely restricted by climatic and edaphic factors, it is vulnerable to extinction. Neither population is on pristine habitat - the larger population is bisected by railroad tracks and a main highway - and both populations have been subject to sheep grazing.
Nation: United States
National Status: N1

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Utah (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (28Sep1978)
U.S. Fish & Wildlife Service Lead Region: R6 - Rocky Mountain

NatureServe Global Conservation Status Factors

Range Extent Comments: Only found in Utah along the Douglas Creek and Gordon Gulch members of the Green River formation in the Wasatch Mountains in Pleasant Valley. This plant was first discovered in 1883.

An intensive search in 1980 by the Utah Native Plant Society (UNPS) revealed a significant number of mature P. ARGILLACEA plants (Smith et al. 1989). This site is located near the Tucker highway rest stop. For purposes of this text, Atwood's find in 1971 will be referred to as the Clear Creek site, and the UNPS-located occurrence will be referred to as the Tucker site, or main site. Collectively, they will be referred to as the Tucker population.

While searching for more P. ARGILLACEA sites in 1989, Ben Franklin (Botanist, Utah Natural Heritage Program) discovered a new population which consists of two subpopulations on open slopes, approximately five miles west-northwest of the Tucker population. For purposes of this text, Franklin's find will be referred to as the Water Hollow-Garner Canyon population.

Clay phacelia occurs in four known sites. Probably only two populations exist, however, due to the close proximity of both pairs of occurrences (Franklin and Tuhy 1989). Harper (1990) suggests that the micro-habitat differences between the Clear Creek and Tucker sites may have effectively "molded" these subpopulations into distinct populations. He believes that the Clear Creek location is both cooler in the winter and warmer in the summer than the Tucker site. His visceral feeling is that the two sites encompass only one population, but he indicates that they may be in the process of diverging.

Number of Occurrences: 1 - 5
Number of Occurrences Comments: Occurs in four known sites; probably only two populations, however, due to the close proximity of both pairs of occurrences.

Population Size Comments: In 1980 Soldier Summit had a population of 200 plants.

Overall Threat Impact Comments: SMALL POPULATION SIZE AND NUMBER: Numerous threats exist for Clay phacelia. Probably the most significant current threat is the species' inherent vulnerability due to small population size and number. In populations with less than 1000 individuals, demographic uncertainties can play a significant role in extinction probability (Shaffer 1987). This may be mitigated by a presumably large soil seed bank. However, the 1987 sheep staging episode significantly impacted the habitat, and the little soil that had accumulated in the area was partly washed away in subsequent thunderstorms. In spite of this, England still suspects a sizeable seed bank (England 1989).

MODIFICATION OR LOSS OF LIMITED HABITAT: Human land-use activities in and near the critical habitats have played the largest role in Clay phacelia's recent dramatic decline. Livestock and sheep have grazed and trampled the plants and their surrounding habitat. The Denver and Rio Grande Western (D&RGW) railroad is adjacent to the Clear Creek site. Construction activities have already modified the neighboring habitat through stabilization of cuts and fills, runoff control, and material storage (Gill et al. 1982). In addition, Highway 6 bisects the Tucker population. Highway maintenance and construction activities such as burning and shoulder stabilization may also impact both potential and existing Clay phacelia and accompanying bee (pollen vector) habitats.

GRAZING BY ENDEMIC SPECIES: Grazing from endemic ungulates and herbivory from small native herbivores has collectively impacted Clay phacelia. Specific threats include mule deer, elk, and occasional moose that frequent the area of critical habitat in the winter. The exposed south- and west-facing slopes provide a welcome winter home with food and relatively warm areas for bedding down. Rock squirrels have also been sighted nibbling on the herbage (Gill et al. 1982).

INTERSPECIFIC COMPETITION: Long-term monitoring of exotic and native species which grow in association with Clay phacelia has not been conducted. However, this should be looked at closely, especially in the absence of grazing. Both MARUBIUM VULGARE and CYNOGLOSSUM OFFICINALE could pose significant competition to P. ARGILLACEA (England 1989).

POTENTIAL THREATS: If pollination is wind-vectored, dust from heavy use of nearby dirt roads could foul the stigma (Harper 1990). The dirt road near the main Tucker site is most frequently traveled during hunting season or late fall. This is well after pollination occurs in Clay phacelia, so it may not be a problem. Large or persistent clouds of fugitive dust have never been observed right near the hillside plants.

Disease or parasitism could also force the extirpation of any of the two remaining populations. Also, a large landslide occurred near Soldier Summit, within miles of the Clay phacelia populations, in the mid-1980s. Obviously, an unforeseen natural disaster of this magnitude could cause extirpation of this taxon from any one of its sites.

Short-term Trend Comments: POPULATION TRENDS: Information regarding population trends is only available for the Clear Creek and Tucker sites (the Tucker population). Since the Water Hollow-Garner Canyon population was discovered in the 1989 field season, there is no available demographic history to date.

TUCKER: In the 1981 growing season, an ocular estimate of 200 to 300 adult flowering plants was made on the Tucker site (England 1989). From 1981 to 1986, similar numbers of adult flowering plants were seen during annual visits to the site. The ratio of rosettes to mature plants was typically 100 to one. During the 1987 growing season, the Tucker population declined drastically. Sheep were bedded down on the slope for two or three days. As a result, the Clay phacelia population was devastated (England 1989). Not one adult plant flowered during the 1987 growing season. England (1989) recalls that the hill "looked like it had been plowed." Return visits to the Tucker site revealed only one rosette in April and May, 6 to 12 rosettes in June, and approximately 100 rosettes in July and August of 1987 (England 1989).

The soil seed bank was severely depleted in 1987 due to the sheep grazing episode and the subsequent erosion from trampling and rainstorms. Since 1987, flowering plants have numbered less than 50. But the ratio of rosettes to flowering adults has remained similar to that which was observed prior to 1987 (England 1989).

CLEAR CREEK: Clear Creek has historically sustained very few Clay phacelia plants. It has ranged from 15 adult plants in 1971 to two in 1979 (Gill et al. 1982), and only held seven mature plants in 1989 (Franklin and Tuhy 1989).

1989 MONITORING OF THE TUCKER POPULATION (BOTH SITES): Concern for this taxon's ability to persist heightened when rosettes, observed in the fall of 1988, were not found the following January at the Tucker site (Franklin and Tuhy 1989). Deer tracks were numerous in the moist soil, and the snow had recently melted. Franklin and Tuhy (1989) therefore surmised that the missing rosettes had probably been grazed by deer. On April 4, 1989, 13 rosettes were located on the Tucker site, presumably only a few weeks old. An additional six rosettes were also found at the Clear Creek site. On May 12, 1989, five more Tucker rosettes were located and one more Clear Creek rosette was found. Numerous seedlings were also present in May, at both the Tucker and Clear Creek sites. Visits to both sites in June 1989 revealed a total of 22 flowering adults.

By August 18, 1989, all of the flowering adult plants were either dead or nearly dead, and numerous rosettes were observed in two locations along the base of an upper cliff-band at the Tucker site. (Franklin and Tuhy 1989). If, in fact, the 1989 adult flowering plants did germinate in the spring of 1989, as observations imply, then Clay phacelia is capable of simulating the life cycle of a summer annual. It was also observed that no new flowering stems had developed on any of the plants since the May 12 visit. In other words, those plants which germinated prior to May 12 were able to mature and reproduce. Those plants which flowered after May 12 remained in vegetative form as basal rosettes (Franklin and Tuhy 1989). This may be a function of moisture availability (Gill et al. 1982).

In the fall of 1989, Lori Armstrong, a Brigham Young University graduate student in botany, began a full-scale research project on the Clay phacelia under the direction of Dr. Kim Harper. To date, 230 Clay phacelia rosettes have been measured, tagged and numbered; 193 at the Tucker site and 37 at the Clear Creek site (Armstrong 1990). More detail regarding the focus of this new study will be discussed in the monitoring and research sections of this abstract.

Intrinsic Vulnerability Comments: Not yet deter. what unique chem. char. of the soil enable this plant to survive at these sites and not elsewhere.

Other NatureServe Conservation Status Information

Distribution
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Global Range: Only found in Utah along the Douglas Creek and Gordon Gulch members of the Green River formation in the Wasatch Mountains in Pleasant Valley. This plant was first discovered in 1883.

An intensive search in 1980 by the Utah Native Plant Society (UNPS) revealed a significant number of mature P. ARGILLACEA plants (Smith et al. 1989). This site is located near the Tucker highway rest stop. For purposes of this text, Atwood's find in 1971 will be referred to as the Clear Creek site, and the UNPS-located occurrence will be referred to as the Tucker site, or main site. Collectively, they will be referred to as the Tucker population.

While searching for more P. ARGILLACEA sites in 1989, Ben Franklin (Botanist, Utah Natural Heritage Program) discovered a new population which consists of two subpopulations on open slopes, approximately five miles west-northwest of the Tucker population. For purposes of this text, Franklin's find will be referred to as the Water Hollow-Garner Canyon population.

Clay phacelia occurs in four known sites. Probably only two populations exist, however, due to the close proximity of both pairs of occurrences (Franklin and Tuhy 1989). Harper (1990) suggests that the micro-habitat differences between the Clear Creek and Tucker sites may have effectively "molded" these subpopulations into distinct populations. He believes that the Clear Creek location is both cooler in the winter and warmer in the summer than the Tucker site. His visceral feeling is that the two sites encompass only one population, but he indicates that they may be in the process of diverging.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map

U.S. & Canada State/Province Distribution
United States UT

Range Map
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U.S. Distribution by County Help
State County Name (FIPS Code)
UT Carbon (49007)*, Utah (49049), Wasatch (49051)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
14 Strawberry (14060004)+*, Price (14060007)+*
16 Spanish Fork (16020202)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A winter annual, 1-4 dm tall, with deeply lobed leaves. The plants produce clusters of lavender-purple flowers in summer.
Technical Description: Clay phacelia reaches 1 to 3.6 dm in height. The stems are finely pubescent, and the pinnatifid leaves, which are oblong in outline, are 0.8 to 5 cm long and 0.5 to 1.4 cm wide (Atwood 1975). The inflorescence is a compound scorpioid cyme, with stipitate-glandular to hirsute pubescence (Gill et al. 1982). The pedicels are 0.7 to 1 mm long, and the cymes elongate to 7.5 cm in fruit (Atwood 1975). The sepals are elliptic to oblanceolate, reaching 2 to 3.8 mm in length and 1 mm in width, with stipitate-glandular pubescence.

The campanulate corolla is bluish-violet, 5 mm in length and width, and its lobes are pubescent. Both stamens and style are exserted (Atwood 1975). The capsule is subglobose, 3.5 mm long and 2.4 mm wide, with glandular and setose pubescence (Gill et al. 1982). The mature seeds are four in number, brown, pitted, ovate to elliptic, 2.4 mm long, and 1.1 mm wide (Atwood 1975). In addition, the seeds have a single groove on one side of the ventral rib (Gill et al. 1982). The ridge is curved, and the seed is more or less excavated along one side (Welsh 1987).

Diagnostic Characteristics: Clay phacelia is very closely related to PHACELIA GLANDULOSA Nutt., a west slope Idaho, Wyoming, and Montana native. In fact, the original collection of clay phacelia in 1883 by Marcus E. Jones, in the vicinity of Pleasant Valley Junction, Utah, was identified P. GLANDULOSA. In 1894, Jones again collected the species; this time at Clear Creek near Soldier Summit in Utah County, Utah. When N.D. Atwood rediscovered the Clear Creek population in 1971, he found it to differ substantially from P. GLANDULOSA. He subsequently described and named it P. ARGILLACEA in 1973 (Gill et al. 1982).

Two plants which grow in association with Clay phacelia, the frequent adventive, CYNOGLOSSUM OFFICINALE (Hound's tongue), and the less frequent native, MENTZELIA LAEVICAULIS, are most readily confused with P. ARGILLACEA. Like Clay phacelia, they both produce basal rosettes. Hound's tongue's leaves, however, are entire and pubescent. MENTZELIA can also be distinguished by its leaves which are lobed with retorse or "velcro-like" hairs (England 1989).

Reproduction Comments: REPRODUCTIVE STRATEGIES: Clay phacelia reproduces sexually. It is unknown whether this species is an obligate selfer, a facultative selfer, or an obligate out-crosser (Harper 1990). England (1989) surmises that apomixis is not part of this taxon's reproductive strategy.

The clay phacelia is considered a winter annual (Atwood 1975), but recent observations suggest that some of the plants in at least one of the extant populations might be simulating the life cycle of a true biennial (England 1989). In addition, observations in the fall of 1988 and the following growing season of 1989 imply that some of the individuals have also simulated the life cycle of a summer annual (Franklin and Tuhy 1989).

Seeds typically germinate in spring and fall, although they sometimes germinate in mid-summer if there is enough available moisture (England 1989). The plant blooms in late July, but non-flowering rosettes have been observed in both June and August (Matheson 1989).

England (1990) speculates that some plants which germinate early in the growing season have insufficient food reserves to initiate flowering the same year. These juvenile or vegetative plants must then over-winter before floral initiation and seed production are possible the following spring. England (1989) verifies that advanced plants have been observed in the spring with early flowering in late May or early June.

Since qualitative monitoring began in the early 1980s, the majority of the Tucker plants have reproduced as winter annuals (England 1989). In this case, the seeds germinate in the fall. Enough biomass is then generated in the fall and subsequent spring, provided they survive the winter, to initiate flowering in late June.

POLLINATION: The pollen vector is unknown; however, a 1988 Bee Biology and Systematics Lab field collection at the Tucker site "yielded the largest series of specimens recorded of a rare species of colletid bee, HYLAEUS GRANULATUS" (Tepedino 1989). Although the very first observed females of this rare bee were among the specimens collected, Harper (1990) surmises that this bee species is far too rare to be the primary pollinator for Clay phacelia. He further speculates that wind, or the more common ground-nesting bees in the area, could prove to be the major pollinators for Clay phacelia. Wind-pollinated species, he adds, tend to have greater genetic diversity.

SEED BIOLOGY/ECOLOGY: As indicated in the technical description, Clay phacelia has the capacity of developing four mature seeds per fruit. In the summer of 1987, England (1989) estimated that as many as 8000 seeds were produced on one adult winter annual. The plant held 200 cymes, with ten flowers per cyme and four seeds per fruit. Needless to say, Clay phacelia can be a very prolific seeder. The seed is produced sequentially over time. Cymes last for about ten days to two weeks. Flowers open daily along the cymes and persist for days before developing into fruit (England 1989).

GENETIC LOAD: Harper (1990) emphasizes the need to gain an understanding of the genetic load ("those genes which are lethal or sublethal in a homozygous state") that this rare plant is carrying. He indicates that studies are necessary to determine the ovule-to-seed ratio within the various populations. This should provide clues to whether the plant is an obligate selfer, facultative selfer or an obligate out-crosser. Obligate selfers, he says, have approximately 90% flower-to-fruit conversion rates, and out-crossed plants have approximately 50% flower-to-fruit conversion rates. By knowing the average number of ovules per flower and the average number of viable seeds produced per number of ovules, one can calculate the approximate number of lethal alleles per ovule, or "conversely determine the efficiency of the pollen vectors" (Harper 1990).

DISPERSAL MECHANISMS: No formal studies have been conducted on dispersal mechanisms. It is surmised by England (1989) that short distant seed dispersal is achieved primarily by wind, water, and gravity. Rosettes are generally located below or in association with mature adult plants. Some long-distance dispersal may be achieved through grazing and defecation, but the plants are probably grazed prior to seed set (England 1989).

Ecology Comments: GERMINATION REQUIREMENTS: Little is known about the germination requirements for Clay phacelia, but steps are being taken to increase our knowledge. Embryo viability of seed collected in June 1988 was tested by means of a tetrazolium (TZ) test. The TZ equaled 95% or 19 viable seeds out of the 20 tested (Matheson 1989). Additional seeds have been collected in June 1989 and on August 21, 1989, but they have not been cleaned or tested yet. Mary Alyce Kobler (Botany, University of Utah), conducted germination studies on PHACELIA LINEARIS and found that seed germination was cued by day length and temperature relationships. She initiated studies on seed germination requirements for P. ARGILLACEA in the fall of 1989 (Matheson 1989). To date seed germination EX SITU has been difficult.

ESTABLISHMENT: Dispersed seeds lodge in cracks or crevices on the shale-covered slopes and germinate when the right (unknown) set of conditions exist. Initial foliage leaves are small, but basal rosettes are formed by early to mid-October. They then over-winter, growing slowly underneath the snow pack. The plants begin to bolt, when the snow melts and the soil temperatures reach sufficient levels in the spring. This usually occurs in May. By late May, the first flowers begin to appear. The plants continually produce more flowers and increase in size, until late June or early July. Prolonged flowering seems to be related to moisture availability. In 1979, flowers were observed as late as mid-October (Gill et al. 1982).

Rosettes which have been dug and removed for EX SITU research and captive propagation in cooperation with the Center for Plant Conservation revealed unexpectedly shallow roots (Matheson 1989). These same plants, which are now in six-inch pots, are being grown in different soil mixes, including native soil, to gain insight into cultural requirements. To date, no significant observations can be reported; however, the rosettes in the vermiculite/perlite soil mix seem to be doing the best (Kobler 1989).

MAINTENANCE: No formal studies have been undertaken to determine basic maintenance requirements for Clay phacelia. However, it is England's (1990) observation that P. ARGILLACEA is dependent upon available, disturbed ground, with little or no competition. He speculates that if the Tucker site were somehow stabilized, P. ARGILLACEA would probably be extirpated. Harper (1990) further theorizes that, if site stabilization did occur, ELYMUS SIMPLEX would tend to dominate, which in the absence of grazing could force extirpation of the species from the Tucker site.

P. ARGILLACEA brings to mind the "chicken or egg" paradox. With the current limitation of knowledge regarding population variances, it is difficult to ascertain the importance of observed population variations in rosette-to-mature-plant ratios (England 1989).

DENSITY AND SUCCESSIONAL STATUS: Extant populations of Clay phacelia cover only three to five percent of the slope's surface area in the patches where they occur. Overall vegetative cover is about ten percent in these same areas of critical habitat (England 1989).

Due to recurrent natural disturbances which erode the surface of the four sites where the plants grow, P. ARGILLACEA can be categorized as a specialized micro-disturbance niche plant (Harper 1990). He notes that some 100 to 125 species have adapted this strategy along the Wasatch Front. The strategy Harper (1990) explains is one of opportunism. Micro-disturbance species are the first to germinate and establish themselves in new areas which have been exposed through some form of natural disturbance (e.g., hooves along the edge of a deer trail or wind and water erosion down a gravelly bank).

HEDYSARUM BOREALE, CLARKIA RHOMBOIDIA, and many PENSTEMONS are examples of plants which are specialized to survive under such conditions. Large disturbances, which are often caused by man's impact on a natural area, can create serious difficulties for some specialized micro-disturbance niche plants. In particular, native annuals frequently "fall apart" on large sites due to increased wind and solar radiation (Harper 1990).

Terrestrial Habitat(s): Forest/Woodland, Savanna, Shrubland/chaparral, Woodland - Conifer
Habitat Comments: BIOLOGICAL HABITAT: Clay phacelia occurs on steep slopes in sparse juniper-pinyon and mountain brush communities (Welsh 1987). The dominant species at the site of the largest element occurrence are RHUS TRILOBATA and AMELANCHIER ALNIFOLIA. At the base of the main slope is a degraded sagebrush steppe. Large and probably influential ungulates are deer, elk, and domestic sheep. Probably the most significant component is mule deer (England 1989). There is mainly wintertime use by big game, and (before a fence was built) transient but heavy spring and fall use by domestic sheep.

Specific plants which grow in the community surrounding the largest Clay phacelia occurrence are: STIPA HYMENOIDES, ELYMUS SIMPLEX, ERIOGONUM BREVICAULE, MENTZELIA LAEVICAULIS, MAHONIA REPENS, OENOTHERA CAESPITOSA, MARRUBIUM VULGARE, CYNOGLOSSUM OFFICINALE, AMELANCHIER ALNIFOLIA, RHUS TRILOBATA, ATRIPLEX CANESCENS, ARTEMESIA TRIDENTATA, CHRYSOTHAMNUS NAUSEOSUS, CERCOCARPUS MONTANUS, ROSA WOODSII, PINUS EDULIS, and JUNIPERUS OSTEOSPERMA. Less frequent plants which also occur in the community are: BROMUS TECTORUM, IVA AXILLARIS, CHENOPODIUM FREMONTII, SYMPHORICARPOS OREOPHILUS, QUERCUS GAMBELLII, CARDUUS NUTANS, AND JUNIPERUS SCOPULORUM (Gill et al. 1982, England 1989, Harper 1990).

PHYSICAL HABITAT: Climate: No long-term weather data exist for the area of the extant populations of this element. Climatological data from a nearby mountain area would represent that of a significantly higher elevation. Similarly, data from a different nearby station would reflect that of a lower valley elevation. The general climate for all of the element occurrence sites can, however, be classified as cool and subhumid (England 1989). A university study (see below) has had instruments nearby.

EXPOSURE: The majority of plants grow on slopes facing west through southeast. The exposure at one site is south and southeast (Gill et al. 1982, Armstrong 1990). The aspect at the other site is also southeast (England 1989). Both sites are free of snow, at least once during the winter, and are typically dry in early spring (Gill et al. 1982). Finally, one subpopulation faces south, and the western subpopulation's aspect is west (Franklin and Tuhy 1989).

SLOPE: At each site, the plants occur on a xeric, exposed slope of the Green River Formation, somewhere between 6000 and 7000 feet elevation (Franklin and Tuhy 1989). The slope at the site which holds the largest number of plants was measured at 70 percent (Armstrong 1990). One occurrence is easily and frequently disturbed by both biotic and abiotic influences. As a result the loose shale on the surface almost continually sloughs down the face of the platey slopes.

SUBSTRATE: The phase of the Green River Formation differs between sites. In each case it is described as a "shaley clay colluvium" (Gill et al. 1982). One occurrence is on a "fine-textured clay derived from a poorly consolidated shale member of the formation" (Gill et al. 1982). The platey shale at one site appears reddish-brown. However, Gill et al. (1982) indicate that the roots of the plants at this site penetrate a buff-to-gray-colored substrate.

One population occurs principally on "a narrow band of fine-textured reddish-brown clay from weathered faces of the Green River Formation" (Franklin and Tuhy 1989). However, further exploration resulted in the location of additional plants growing in a "grey-white, small fragmented shale occurring above the reddish-brown clay layer" (Franklin and Tuhy 1989).

Analysis of a soil sample from one site revealed a pH greater than eight. Further, greenhouse cultivation of the seedlings in native and artificial soils indicates that the clay composition of the native soil enhances its ability to hold moisture, even under conditions of high solar radiation (Kobler 1989).

Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: In summary, stewardship priorities for the Clay phacelia are as follows:

1. Protect all element occurrences, their pollinators, and their habitat's from grazing, trampling, burning, or any other potential or immediate threats. Maintain the game fence surrounding the Tucker through bimonthly checks, and cage as many additional plants at other sites as is feasible.

2. Conduct monitoring and research studies. Understand the life history of this taxon well enough to recommend and predict results from active management.

3. Determine the relative genetic diversity within individual element occurrences, as well as for the whole extant gene pool.

4. Determine methods for producing genetically diverse seed and plants, EX SITU, for potential introduction and reintroduction, as well as germplasm preservation.

5. Search for more element occurrences and identify viable introduction sites. Determine whether efforts to establish new methods were successful.

6. Monitor the Tucker site using semi-resident volunteer help.

7. Initiate and complete other actions called for in the final recovery plan approved and issued by the Fish and Wildlife Service.

Restoration Potential: The Tucker population seems to be recovering from the 1987 sheep staging episode. In spite of the significant decline in the numbers of plants, individual plant vigor is currently similar to the plants observed in the early 1980s (England 1989). Clay phacelia seemingly has some inherent resilience.

Harper (1990) further notes that if Clay phacelia were a true biennial, its chances of survival would be less than that of an annual. A biennial needs two complacent growing seasons back-to-back, in order to successfully complete one life cycle. Annuals only need one.

The Nature Conservancy erected a game fence (July 1990) for the elimination of all ungulate species at the Tucker site. This will give Clay phacelia a viable chance to recover on this site, without the tremendous grazing and trampling pressures it has experienced in recent years. Inter-specific competition, particularly that from exotic species, will need to be closely monitored in the absence of grazing. Ultimately, it remains to be seen whether P. ARGILLACEA is resilient enough to sustain itself without a fence to protect it from the indigenous ungulates. Surely, it will always need some degree of protection from livestock.

Preserve Selection & Design Considerations: NATURE OF HABITAT: Limited habitat is one of the major constraints for the perpetuation of this species. Clay phacelia "grows on a sequence of strata in the Garden Gulch and Douglas Creek members of the Green River Formation. These members are of limited distribution in the portion of Utah were there might be dispersal of the species. Still, the substrate is of sufficient breadth that very large populations could presumably be supported if there were no other limiting factors on the species" (Gill et al. 1982).

For an in-depth look at potential introductory sites refer to the Franklin and Tuhy Report (Franklin and Tuhy 1989), Appendix 1. In this report, they discuss areas which have been extensively searched for Clay phacelia. They also recommend specific, future searches for the taxon, and even re-searches in wetter years.

BUFFERS AND MITIGATION OF OFF-SITE THREATS Construction activities along the railroad near the Clear Creek population have significantly altered Clay phacelia's habitat. Gill et al. (1982) indicate that the D&RGW railroad personnel are aware of the plant's status and that they seemingly have a positive attitude toward its needed preservation. Regardless, open annual communication is probably a wise, minimum, preventative measure to keep the company's awareness level up.

Close monitoring of activities involving railroad or highway construction is also recommended. Ground nesting bees dislike disturbance due to vegetative clearing along ditch banks, burning, fence construction, etc. If they turn out to be the main pollen vector for Clay phacelia, it will be prudent to monitor and perhaps negotiate modified land management procedures in the general area, which could disturb the bees habitat (Harper 1990).

Management Requirements: The clay phacelia is "one of the most important plants to protect" in Utah. It was the sole purpose of a protracted land acquisition effort that led to establishment of the Soldier Summit Preserve in 1991. The preserve contains about 70 acres, is located in north-central Utah, and is managed by the Great Basin Field Office (GBFO) of The Nature Conservancy. The species' federal status is "Endangered" and its natural heritage program ranking is G1S1.

Larry England (1990) of the USFWS describes protection of the Tucker site as being "paramount to the perpetuation of this species." Preservation of the site and protection of the plants from ungulate trampling and grazing is probably the highest management priority. Secondarily, stewards must advance understanding the biology and ecology of this element well enough to enhance it through management on existing sites and for the selection of viable introduction sites. In the meantime, active protection should be sought for all element occurrences.

IN SITU MANAGEMENT The largest element occurrence is under the control of The Nature Conservancy, Great Basin Field Office. Maintenance of a seven-foot-high game fence surrounding the 70-acre preserve will help immensely. Although fencing is not likely on all four occurrence sites, protective easements should be pursued and negotiated with respective land owners. Management procedures should focus on caging or exclosure methods to provide effective grazing protection for as many individual Clay phacelia plants as possible.

Field management should also include the monitoring of any D&RGW and highway management activities which could harm the element occurrences both on and off the nature preserve.

EX-SITU MANAGEMENT Captive seed propagation and seed production are key activities which will help ease the seemingly high probability of extinction for this rare taxon. If demographic, environmental, or catastrophic events were to eliminate the species, the captive material would be the only remaining germ plasm.

In addition, if seeds and plants can be successfully propagated, they could be essential in future introduction or reintroduction attempts on selected sites.

Monitoring Requirements: Harvey (1985) suggests that demographic studies of rare plants provide valuable information on life cycle features which could lead to the identification of critical stages in an element's life cycle which contribute to small population size.

Population trends for the new Water Hollow-Garner Canyon population are unknown. The Clear Creek and Tucker sites have been observed long enough to confirm that there has been a long-term decrease in plant numbers at both locations. In addition, the population at the main site has decreased in the mid-term. There has been a short-term increase in plants since the 1987 decline, and this has probably been accentuated by emplacement of the fence. Although overall trends are apparent, monitoring of individual plants is vital to the understanding of the life history of this taxon.

Harper (1990) encourages monitoring of all sites because micro-habitat peculiarities may be determining unique genotypes. As mentioned earlier, in situ studies should emphasize low impact methods, since significant substrate disturbance can result with every footstep on the steep, shale-covered slopes.

FIELD DATA COLLECTION: Menges (1986) recommends that field data collection on mortality, growth, and reproduction of individual plants should be conducted over time. He suggests that measurements might include reproductive status, number of stems, stem diameter, height, number of leaves, and leaf and rosette sizes. Individuals should be grouped into stages (e.g., seedling, juvenile, vegetative, or reproductive plants), as opposed to attempting to determine specific ages. Plants should be marked using any variety of standard methods and mapped in detail.

MONITORING DURATION: Menges (1986) indicates that two field seasons are generally enough to make preliminary analyses; however, he hastens to point out that predictions are "only as good as that time period is typical of the future." The current stochastic (uncertainty) approaches to population modeling require, according to Menges (1986), "enough measurements to estimate both temporal and spatial variability in life history components." In other words, comprehensive, long-term monitoring is essential for demographic analysis and for effectively predicting the extinction probability and minimum viable population for a given element. See Menges (1986) for discussion on matrix projections, transient predictions, and life history analysis.

RECENT FINDINGS: Menges (1986) summarizes some recent findings in modeling plant species. He indicates that modeling efforts can be based on modest levels of data collection; that long-term modeling is desirable in light of the significant impact environmental fluctuations have on slow-growing perennials (not applicable to Clay phacelia but could be important with respect to inter-specific competition); that the greatest amount of monitoring effort should prioritize the fate of established plants; and that quantifying environmental variability is of the utmost importance for effectively predicting the future of rare plants.


Management Programs: IN SITU MANAGEMENT PROGRAMS: Cooperation between the Nature Conservancy, USFWS, Utah Natural Heritage Program, and the BYU botanists will result in the placement and monitoring of cages at the other element occurrence sites (besides the Tucker site).

EX SITU MANAGEMENT PROGRAMS: The RBG&A Horticulture Director and Ms. Kobler, in cooperation with the CPC, are intending to collect seed from multiple sites. This will ensure genetic diversity in the captively propagated plants and the seed bank for protection of the gene pool. Researched cultural techniques for domestic seed and plant production of Clay phacelia may then be implemented when introductory or reintroductory efforts begin.

For further information on propagation programs, contact:

Larry England, Botanist, US Fish and Wildlife Service, 1745 W 1700 S, Room 2078, Salt Lake City, UT 84104, (801) 524-4330.

Monitoring Programs: 1989 FIELD SEASON: Monitoring of the Clay phacelia began in early spring of 1989 by individuals from the Utah Natural Heritage Program, under contract with the Uinta National Forest. The primary goal of the agreement was to search for additional P. ARGILLACEA occurrences within the Uinta National Forest. However, a secondary goal was to protect and monitor the growth of individual Clay phacelia plants on both the Clear Creek and Tucker sites.

Site visits occurred on January 31, April 4, May 12, June 15, July 13, and August 18, 1989. Rosettes were located each visit, and the diameter was measured both up and down the slope, as well as across. When the plants began to bolt, their height was measured. The resultant number of flowers produced per individual plant was also estimated. Protective cages were placed over as many rosettes in early spring as was possible. For detailed notes on these monitoring efforts refer to the Report for 1989 Challenge Cost Share Project, Uinta National Forest, prepared by Franklin and Tuhy (1989).

A second monitoring effort began this fall. Dr. Kim Harper and a graduate student, Lori Armstrong, have run a zig-zagged 300-meter tether rope up the slope, through the largest occurrence of P. ARGILLACEA at the Tucker site. Galvanized nails, with numbered aluminum tags, have been driven into the substrate, adjacent to every plant within one meter of the rope. The diameter of each tagged rosette has been noted. In addition, all of the plants which are growing underneath the protective cages throughout the site have also been measured and numbered. To date, 193 plants have been measured and marked at the Tucker site. Across the highway, at the Clear Creek site, 37 plants have also been tagged and measured. Protective cages are in place at this site, as well (Harper and Armstrong 1989).

Current monitoring plans include: population studies following the fate of 300 plants (mortality by period); reproductive studies determining the number of flowers per plant, fruits per plant, seeds per fruit, and ovules per mature seed; and pollination studies estimating the genetic load inherent in this rare plant.

Inter-specific competition will also be studied by monitoring vegetation within a permanently marked inventory plot. This will create a linear record on all plants within the plot. Distance relationships to the four nearest neighbors of some individually tagged Clay phacelia plants will be plotted against the latter's terminal size and seed set. Regression analysis, using distance to the nearest plant in relationship to size and vigor of the correlating Clay phacelia plants, will provide valuable insight into the dynamics of inter-specific competition (Harper 1990). The physical climate will also be monitored through placement of a weather station on Grant William's land across the road from the main site. Duration for the monitoring study will be a minimum of two years. Further study is subject to funding availability (Harper 1990).

For further information on these programs, contact:

Dr. Kimball Harper,Department of Botany, Brigham Young University,Provo, UT 84602, (801) 378-2129

Mr. Larry England, Botanist US Fish and Wildlife Service, 1745 W 1700 S, Room 2078, Salt Lake City, UT 84104, (801) 524-4430

Mr. Ben Franklin, Botanist Utah Natural Heritage Program, 3 Triad Center, Suite 400, Salt Lake City, UT 84180-1204, (801) 538-5524

Management Research Programs: GERMINATION REQUIREMENTS: Mary Alyce Kobler (Botanist, University of Utah), in cooperation with The Red Butte Gardens & Arboretum, University of Utah (RBG&A), and The Center for Plant Conservation (CPC), is currently conducting studies on P. ARGILLACEA germination requirements. She is studying the effects of temperature fluctuations, light and dark treatments, and soil manipulation, EX SITU.

CULTURAL REQUIREMENTS: In addition to seed germination work, Ms. Kobler is hoping to produce enough plants to study cultural requirements and seed crop production techniques.

REPRODUCTIVE BIOLOGY: It has been suggested that pollination studies be carried out EX SITU at Utah State University, by associates from the Bee Biology and Systematics Lab. Harper and Armstrong are also planning to carry out some pollination and seed dispersal studies, IN SITU, during the 1990 field season (Harper 1990).

GENETIC DIVERSITY: Harper (1990) states that seed proteins evolve at a faster rate than leaf proteins. In addition to the above-described monitoring work, Harper and Armstrong hope to conduct some isozyme studies on seed samples from each of the element occurrence sites. Dan Fairbanks (Assistant Professor of Botany, BYU) has developed good micro-technology techniques for such research and will likely work with them on this project (Harper 1990).

For more information on research programs, contact:

Dr. Kimball Harper,Department of Botany, Brigham Young University, Provo, Utah 84602 (801) 378-2129

Lori Armstrong 1120 E 2680 N, Provo, Utah 84604, (801) 378-4955 (W), (801) 377-4664 (H)

Mary Alyce Kobler, Greenhouse Manager, 406 S. Biology, University of Utah, Salt Lake City, Utah 84112

Mary Pat Matheson, Director of Horticulture, Red Butte Gardens & Arboretum Building 436, Room 116, University of Utah, Salt Lake City, Utah 84112

Vincent J. Tepedino or Terry Griswold, Research Entomologist, Bee Biology & Systematics Lab, Utah State University, Logan, Utah 84322-5310, (801) 750-2559 or 750-2526

Management Research Needs: BIOLOGY/ECOLOGY: Long-term monitoring data will aid significantly in correlating life cycle variability to environmental fluctuations and inter-specific population dynamics. Understanding the basic biology and ecology of this rare plant will enhance our ability to effectively protect and increase existing and future populations. For instance, it would be very helpful to understand the essential habitat requirements inherent to this taxon.

Studies should focus on substrate differences between existing populations. To what degree do they differ? What are the water, light, nutritional, propagation, and growth requirements for P. ARGILLACEA? Which animals and insects feed on the herbage or other plant parts? What are Clay phacelia's outside tolerances with respect to inter-specific competition? This sort of information should be pursued, as it will be invaluable for making wise management related decisions on existing sites. In addition, it will allow for intelligent identification and selection of viable introductory sites.

MONITORING AND RESEARCH APPLICATION: Monitoring data coupled with information from research studies on the general biology of this taxon will aid in the development of a projection matrix. An overriding question remains unanswered: Why has the Clay phacelia always had such a narrow distribution over the last century? Matrix projection techniques may be helpful in determining the constraining factor(s). For more information regarding this concept, see Menges (1986).

Population/Occurrence Delineation
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Minimum Criteria for an Occurrence: A natural occurrence of one or more plants.
Separation Barriers: EOs are separated by either: 1 kilometer or more across unsuitable habitat or altered and unsuitable areas; or 2 kilometers or more across apparently suitable habitat not known to be occupied.
Separation Distance for Unsuitable Habitat: 1 km
Separation Distance for Suitable Habitat: 2 km
Separation Justification: The rationale for this large a separation distance across suitable but apparently unoccupied habitat is that it is likely additional research will find this habitat to be occupied. It can often be assumed that apparently unconnected occurrences will eventually be found to be more closely connected. No information on mobility of pollen and propagules is available on which to base the separation distance for this species.
Date: 21Jun2002
Author: Ben Franklin
Population/Occurrence Viability
Help
Excellent Viability: SIZE: 500 or more individuals (based on available EOR data). CONDITION: The occurrence has an excellent likelihood of long-term viability as evidenced by the presence of multiple age classes and evidence of flowering and fruiting, indicating that the reproductive mechanisms are intact. This occurrence should be in a high-quality site with less than 1% cover of exotic plant species and/or no significant anthropogenic disturbance. LANDSCAPE CONTEXT: The occurrence is surrounded by an area that is unfragmented and includes the ecological processes needed to sustain this species. This includes the presence of the appropriate edaphic requirements of this species, i.e., in a matrix of fine textured soils and fragmented shale of the Green River Formation, in a matrix of pinyon-juniper woodlands.
Good Viability: SIZE: 200 to 499 individuals (based on available EOR data). CONDITION: 200 to 499 individuals (based on available EOR data). LANDSCAPE CONTEXT: The occurrence should have a good likelihood of long-term viability as evidenced by the presence of multiple age classes and evidence of flowering and fruiting, indicating that the reproductive mechanisms are intact. Anthropogenic disturbance within the occurrence is minimal. If exotic species are present, they comprise less than 10% of the total ground cover.
Fair Viability: SIZE: The surrounding landscape should contain the ecological processes needed to sustain the occurrence but may be fragmented and/or impacted by humans. CONDITION: 20 to 199 individuals (based on available EOR data). LANDSCAPE CONTEXT: The occurrence may be less productive than the above situations, but is still viable, with multiple age classes and evidence of flowering and fruiting, indicating that the reproductive mechanisms are intact. The occupied habitat is somewhat degraded (exotic plant species make up between 10-50% of the total ground cover and/or there is a moderate level of anthropogenic disturbance).
Poor Viability: SIZE: There may be significant human disturbance, but the ecological processes needed to sustain the species are still intact. CONDITION: Less than 20 individuals (based on available EOR data). LANDSCAPE CONTEXT: Little or no evidence of successful reproduction is observed (poor seedling recruitment, no flowering or fruiting observed, or poor age class distribution). Exotic plant species make up greater than 50% of the total ground cover, and/or there is a significant level of human disturbance.
Justification: SIZE: Large populations in high quality sites are presumed to contain a high degree of genetic variability, to have a low susceptibility to the effects of inbreeding depression, and to be relatively resilient. EOs not meeting "C"-rank criteria are likely to have a very high probability of inbreeding depression and extirpation due to natural stochastic processes and/or occur in degraded habitat with low long-term potential for survival. CONDITION: Large populations in high quality sites are presumed to contain a high degree of genetic variability, to have a low susceptibility to the effects of inbreeding depression, and to be relatively resilient. EOs not meeting "C"-rank criteria are likely to have a very high probability of inbreeding depression and extirpation due to natural stochastic processes and/or occur in degraded habitat with low long-term potential for survival. LANDSCAPE CONTEXT: Large populations in high quality sites are presumed to contain a high degree of genetic variability, to have a low susceptibility to the effects of inbreeding depression, and to be relatively resilient. EOs not meeting "C"-rank criteria are likely to have a very high probability of inbreeding depression and extirpation due to natural stochastic processes and/or occur in degraded habitat with low long-term potential for survival.
Key for Ranking Species Element Occurrences Using the Generic Approach (2008).
U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
Help
Authors/Contributors
Help
NatureServe Conservation Status Factors Edition Date: 10Sep1986
NatureServe Conservation Status Factors Author: Greene, L., rev. D. Atwood, rev. B. Franklin (1996)
Management Information Edition Date: 28Jan1990
Management Information Edition Author: DEBORAH COX CALLISTER; NICHOLAS VAN PELT

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
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  • 100th Congress. 1988. Endangered Species Act of 1973, appropriations authorization for fiscal years 1988-1992. Public Law 100-478-October 7, 1988 102 STAT.2307-102 STAT.2323.

  • Armstrong, L. 1990. "PHACELIA ARGILLACEA 1990-1992 Study Objectives". Unpublished typescript, 3 pages.

  • Armstrong, L. 1990. Map of new Phacelia argillacea locations at EO #003.

  • Armstrong, V. A. 1992. Site characteristics and habitat requirements of the endangered clay phacelia (Phacelia argillacea Atwood, Hydrophyllaceae) and Aspects of the biology of Phacelia argillacea Atwood (Hydrophyllaceae), an annual. Manuscripts of two journal articles presented to the Department of Botany and Range Science[,] Brigham Young University[, Provo, Utah]. In partial fulfillment of the requirements for the degree Master of Science. 65 pp.

  • Atwood, N. D. 1973. Two new species of Phacelia (Hydrophyllaceae). Phytologia 26(6): 437-438.

  • Atwood, N. D. 1978. Status report for Phacelia argillacea. Unpublished report prepared for the U.S. Fish and Wildlife Service. 6 pp. [updated April 10, 1980]

  • Atwood, N.D. 1975a. A revision of the Phacelia Crenulatae group (Hydrophyllaceae) for North America. The Great Basin Naturalist 35(2): 127-244.

  • Bean, P. 1997. Botanists take big step to save tiny plant. Standard-Examiner [June 23]. Pp. 1B, 4B.

  • Callister, D.C. and N.V. Pelt. 1992. Element Stewardship Abstract for PHACELIA ARGILLACEA[,] clay phacelia. The Nature Conservancy. 21 pp.

  • Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. L. Reveal and P. K. Holmgren. 1984. Intermountain Flora: Vascular Plants of the Intermountain West, U.S.A., Volume 4. Bronx: New York Botanical Garden. 573 pp.

  • Cronquist, A., A.H. Holmgren, N.H. Holmgren, J.L. Reveal, and P.K. Holmgren. 1984. Intermountain Flora: Vascular Plants of the Intermountain West, U.S.A. Vol. 4, Subclass Asteridae (except Asteraceae). New York Botanical Garden, Bronx. 573 pp.

  • Franklin, M.A., and J.S. Tuhy. 1989. Report for 1989 Challenge Cost Share Project, Uinta National Forest. Target species: Phacelia argillacea Atwood (clay phacelia). Utah Natural Heritage Program, Salt Lake City. 7 pp + appendices.

  • Gill, J., D. Atwood, A.W. Heggen, J.L. Miller, Sr., S.L. Welsh, and J. Palmer. 1982. Phacelia argillacea Atwood recovery plan. U.S. Fish and Wildlife Service, Denver, Colorado. 13 pp.

  • Gill, J., D. Atwood, A.W. Heggen, J.L. Miller, Sr., S.L. Welsh, and J. Palmer. 1982. Phacelia argillacea Atwood recovery plan. U.S. Fish and Wildlife Service, Denver, Colorado. 13 pp.

  • Harper, K., and L. Armstrong. 1992. Report for 1991 Challenge Cost Share project, Uinta National Forest. Target species: Phacelia argillacea Atwood (clay phacelia). Prepared for USDI Bureau of Land Management, Utah State Office, Salt Lake City. 14 pp. + appendices.

  • Harvey, H.J. 1985. Population biology and the conservation of rare species. In J. White. Studies on plant demography. John L. Harper, Academic Press. Orlando, FL. pages 111-123.

  • Herbarium of Brigham Young University. Photocopies of herbarium specimen for PHACELIA ARGILLACEA.

  • Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.

  • Menges, E.S. 1986. Predicting the future of rare plant populations: Demographic monitoring and modeling. Natural Areas Journal 6: 13-25.

  • Montague, C. 1990. Letters with reference to activities concerning Phacelia argillacea.

  • Nielsen-Snyder, K. 1984. Preliminary inventory for endangered and thereatened species for the Utah County Planning Unit, Pony Express Resource Area.

  • Shaffer, M. 1987. Minimum viable populations: Coping with uncertainty. In M.E. Soule (ed.). Viable populations for conservation. Cambridge University Press, Cambridge, Great Britain.

  • Smith, F.J., K.D. Heil, and J.M. Porter. 1989. Clay phacelia, PHACELIA ARGILLACEA Atwood recovery plan. Unpublished report prepared under contract with USFWS. Denver, CO. 17 pp.

  • Soper, C. 1987. Utah Preserve Design. 7 pp. + two 8-1/2 x 11 maps.

  • Tepedino, V.J. 1989. Bee Biology & Systematics Lab, Utah State University. Unpublished letter to Chris Montague of The Nature Conservancy, Salt Lake City, UT. Mar 27, 1989.

  • The Nature Conservancy. 1990. Element stewardship abstract (ESA) on RANUNCULUS ACRIFORMIS var. AESTIVALIS. Unpublished report prepared by D.C. Callister under contract with The Nature Conservancy, Salt Lake City, UT. 23 pp.

  • U.S. Fish and Wildlife Service. 1978. Determination of five plants as endangered species. Federal Register 43(189): 44810-44812.

  • Utah Native Plant Society. 1980. Utah Native Plant Society: field survey of potential habitat areas for clay phacelia (PHACELIA ARGILLACEA). 5 pp. + map.

  • Welsh, S. L. 1978. Endangered and threatened plants of Utah: a reevaluation. Great Basin Naturalist 38(1): 1-18.

  • Welsh, S. L., N. D. Atwood, S. Goodrich, and L. C. Higgins [eds]. 1993. A Utah Flora (2nd ed., revised). Provo, UT: Brigham Young University. 986 pp.

  • Welsh, S.L. 1979. Illustrated manual of proposed endangered and threatened plants of Utah. Brigham Young Univ., Provo, UT. 318 pp.

  • Welsh, S.L., N.D. Atwood, L.C. Higgins, and S. Goodrich, eds. 1987. A Utah Flora. Great Basin Naturalist Memoir 9, Brigham Young University, Provo, Utah. 894 pp.

  • Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Higgins (eds.) 1993. A Utah flora. 2nd edition. Brigham Young Univ., Provo, Utah. 986 pp.

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