Transition Monitoring for

Riparian Vegetation in the Colorado River Corridor of

Grand Canyon National Park

Michael J.C. Kearsley

and

Tina J. Ayers

Department of Biological Sciences Northern Arizona University Flagstaff, AZ 86011

Starting Date: 1 October 1995

Duration: 12 Months

Submitted To: U.S. Department of the Interior Bureau of Reekunation Glen Canyon Environmental Studies Flagstaff, Az 86002-2459 ‘1 24

MI D. Henry 0. Hooper, Ph.D. Co-Princip Investigator Associate Vice President for Academic Affairs, - 4 Research and Graduate Tina .7. Ayers, StucUes Co-Principal Investigator ABSTRACT

Riparian in marsh, channel margin, and back beach environments downstream of Glen Canyon Dam are strongly affected by the timing, magnitude, and duration of water releases. During the Bureau of Reclamation's Interim Flows, the number of channel margin marshes has increased, but their total area has decreased. Also, there has been a riverward displacement of wet marsh (Typha, Juncus, Phragmites) species in return current channel marshes. Water stress in coyote willow, Salix exigua, and dieback of cattails, Typha domingensis, are inversely related to discharge volumes.

Here we propose monitoring of riparian and wetland assemblages using aerial photographs and ground-truthing as outlined in the Transition Monitoring Plan strategy document. We will track the development of these assemblages, paying special attention to fluvial marshes and newly exposed habitats in the zone between 20,000 cfs stage elevation and 28,000 cfs stage elevation (the 'New Dry Zone'). We will use mapping and census methods developed and refmed in the last two years of Interim Flows monitoring. In addition to providing broad scale patterns of vegetation change, the data will reveal patterns of species loss and turnover and the establishment and spread of important exotic plant species

This information is important for both land- and water- management agencies because riparian plant assemblages are continuing to change rapidly under Interim Flows. For example, the higher elevation areas of marshes lost significant numbers of obligate wetland species during the first three years of Interim Flows. These same species colonized the New Dry zone during that same time. Also, riparian and wetland vegetation have special significance for wildlife in and near the river corridor.

)(earsley and Ayers 2 INTRODUCTION

Problem Statement The timing, magnitude, and duration of water releases from Glen Canyon Dam have had strong effects on new high water zone plant assemblages in the river corridor of Glen Canyon National Recreation Area and Grand Canyon National Park (Campbell and Green 1968, Carothers and Aitchison 1976, Phillips et al 1977, Johnson et al. 1985, Koslowski 1984, Stevens et al. 1995). Through physical effects of substrate deposition and scouring and indirect, groundwater-mediated influences, dam operations can determine the identity, diversity, and productivity of riparian plant assemblages (Stevens and Ayers 1992, Stevens 1989, Turner and Karpisak 1980). These vegetation characters, in turn, determine the suitability of these settings for wildlife.

Given these effects, vegetation monitoring must continue, even at a minimal level. Although Interim Flows (IF) discharge criteria were mandated at Glen Canyon Dam to minimize the effects on downstream resources, riparian habitats have continued to change rapidly since their implementation. Fluvial marshes have become drier, measured either by changes in assemblage-wide ordinations or basal area of obligate wetland species (Kearsley and Ayers 1995). Habitats in the 'New Dry' zone (between the maximum stage discharge elevations of Interim Flows and pre-IF normal flows) have been colonized rapidly by a diverse group of species.

Specific Objectives We propose to continue monitoring the composition and successional trajectories of plant assemblages in the river corridor. The data will describe plant assemblages and will provide updates on exotic plant species. We cannot overemphasize too strongly the importance of data continuity to the monitoring effort, given the dynamic nature of riparian plant assemblages and the importance of riparian vegetation to other species in the river corridor. Our work will be directed towards answering the following questions:

1. Are riparian plant assemblages below the old high water zone continuing to change during the extension of interim flows? This question breaks down to specific questions: a). Are there losses of species or a change in the abundance of important wetland and riparian species in our study areas? We will be testing the null hypothesis of no change in either species composition or species abundance. b). Are the areas in which obligate wetland species occur continuing to shrink or changing position as a result of water release patterns? Our null hypothesis is that the areal extent and location of areas in which these species will not change. c). Are populations of exotic species growing and expanding into new areas during the study period? We will test hypotheses of no changes in foliar cover of specific exotic species in our study sites and no movement into new areas in the study sites.

2. Have the previous year's flows affected the physical layout of return-current channel marshes in our study sites? We will combine our efforts with the NAU sandbar studies

Kearsley and Ayers 3 group to test the hypothesis that the morphologies of return current channels have been unaffected by flows during water year 1995.

In addition, we will be refilling the integration of previous vegetation studies' data into the GCES GIS. Although most data from 1992 onward were collected in ways which allow them to be integrated in a very gross manner (i.e. references to data files), a better integrated set of data would allow easier access to the information and better use of the capabilities of the GIS.

Process of Integration with Transition Monitoring Plan. The work outlined in questions 1 through lc above directly address Item IV: A(1) in the Transition Monitoring Plan. Question 2 above addresses Item IV: A(2)

BACKGROUND

Review of Literature and Previous Work Turner and ICarpisak (1980) and Pucherelli (1988) reported a rapid increase in riparian vegetation cover in the new (post-dam) high water zone (NHWZ) between 1965 and 1982. Cover was significantly reduced in this new riparian zone in wide reaches during spillover flooding in 1983-1984. Brian (1982, 1988) and Stevens and Waring (1988) reported significant loss of riparian vegetation in the NHWZ as a result of spillover flooding. The latter authors noted that spillover flooding degraded substrate, decreased riparian plant cover and diversity, and stimulated germination.

Stevens (1989) compared the impacts of flooding on riparian vegetation in tributaries and along the river corridor. He concluded that dam-induced substrate changes in soil texture, water availability, and nutrient status were largely responsible for the successional development of the phreatophyte community. He also reported that the river corridor was structurally more similar to the sparsely-vegetated ephemeral tributaries than to the richly- vegetated perennial tributaries. Stevens and Waring (1988) studied post-1983 development of woody riparian vegetation along the Colorado River in Grand Canyon National Park from 1984 through 1988, reporting that germination of dominant riparian species was induced by flooding, and exotic Tamarix ramosissima density appeared to have increased as a result of dam operations.

GCES Phase II studies on riparian vegetation were conducted by Ayers and co- workers (Stevens and Ayers 1993, Stevens et al. 1995). Their data defmed several impacts of discharge on riparian vegetation relevant to this proposal:

1. Reduced flooding frequency after completion of Glen Canyon Dam in 1963 allowed fluvial marshes to form in low velocity channel margin environments in wide reaches.

Kearsley and Ayers 4 Marshes were scoured by spillover flooding in 1983 and have redeveloped since 1987. Approximately 253 wet and 850 dry marshes were found in the Colorado River corridor in the park in 1991. Total marsh area was conservatively estimated at 25.0 ha, with 30% as wet marsh (Typha domingensis, Juncus spp. and Scimus spp) and 70% as dry marsh (Phragmites australis and Equisetum X ferrissiz). Marshes in this system were generally small (< 0.5 ha), with wet marshes significantly larger than dry marshes. Marsh assemblage composition and succession was related to sediment distribution and daily inundation frequency (0.4 to 0.9 for wet marshes). More than 95% of the wet marsh vegetation in this system lay between the 283 m3/sec (10,000 cfs) and 566 m3/sec (20,000 cfs) stages. Fluvial marshes can re-develop rather quickly after flooding events (e.g. 1983-1986).

2. Riparian seedling establishment was greatest in the New High Water Zone and was dominated by herbaceous species there. Seedling density varied as a function of particle size, and differed between plant groups (xerophytes and phreatophytes) in wide versus narrow channels. Tamarix ramosissima seedling density was low in 1991, but limited establishment suggests that this species is capable of continuing to colonize the river corridor under fluctuating flow regimes.

3. Several previously unrecorded non-native species colonized the river corridor. Erianthus ravennae (a large Eurasian bunchgrass) and Lepidium latifolium, a clonal herb had rapidly colonized riparian habitats in the river corridor. Prompt action was requested to prevent these species from successfully invading the river corridor.

After Phase II studies, vegetation monitoring continued during Interim Flows (Stevens and Ayers 1995, Kearsley and Ayers 1995). The major findings and products from these studies are listed below:

1. We developed a workable vegetation ordination system for quantifying trajectories of plant assemblages through time. First, marsh census data and topographic survey information have been combined to examine the successional trajectories of marsh plots as a function of dam discharge patterns. For each of 7 marshes, census data from spring and fall for the two years of Interim Flows have been analyzed separately, owing to large and significant differences in the populations of annual plant species in the two seasons. Using detrended correspondence analysis (a principle components-style analysis with allowances for strong environmental gradients), each plot was given a score for each year on the first canonical axis, which represented moisture availability (Kearsley and Ayers 1995, Kearsley et al. in prep). Repeated measures analysis of variance on these data show that both spring and fall censuses had more dry-site species, fewer wet-site species and more individuals of dry-site species in the second year of censusing.

2. We have developed a mapping and vegetation classification scheme which integrates well with the GCES-GIS. Using maps derived from aerial photos, ground truthing, and

Kearsley and Ayers 5 two-way indicator species analysis, we have mapped and classified broad-scale vegetation patterns. Twenty-six major vegetation types (sensu Brown et al. 1979) have been distinguished and their distribution and extent mapped in 6 of the GCES- GIS reaches. We expect this method will become a monitoring tool of increasing importance as refinements are made to the GIS and its interface with the vegetation data. We have also used the same kind of classification scheme with the long-term quadrat data, but it doesn't lend itself to inclusion in the GIS as well as the mapping data.

3. The number of patches of marsh vegetation is increasing between Lee's Ferry and Diamond Creek, but the total area of marsh vegetation in that reach has decreased in the past two years. In 1993 we found 895 cattail/rush/reed fluvial marsh patches (7.03 ha) between Lees Ferry and Diamond Creek. This represented 3.5 fold increase in cattail/rush/reed marsh patch density, up from 253 in 1991, but a 22 percent decrease in marsh area, down from 9.0 ha in 1991. The great increase in cattail/rush/reed marsh density/km reflects the large number of small, wet marsh patches that are presently colonizing the river corridor. Despite this large increase in density, total marsh area decreased between inventories. The reaches downstream from the Little Colorado River that had been inundated and scoured by the January- February 1993 floods decreased in area most strongly, with marsh area losses of up to 90 percent in some narrow reaches. In contrast, the four unflooded reaches upstream from the Little Colorado River generally exhibited a slight gain in marsh area.

4. Recently exposed habitats between 225 and 750 m3/sec (20,000 and 30,000 cfs) has been rapidly colonized under Interim Flows. The new dry zone quadrats established in our study areas were devoid of vegetation at the inception of Interim Flows, as determined from close-level aerial photographs taken in mid-summer, 1991. Quadrat analyses in 1992 suggested that the new dry zone was extremely productive, and had become almost as densely colonized as the bar platform (general beach) zone, with little detectable difference in basal area of plant life or stem density between the two zones (F1,21 = 2.979, p = 0.099; total basal area data were arcsine squareroot transformed). The new dry zone exhibited comparable basal areas of all assemblages except woody non-clonal phreatophytes when compared to the bar platform zone. The bar platform and new dry zones support only about 50 percent of the species richness and basal area of the other geomorphic settings under study, a finding in accord with the conclusions of Stevens and Ayers (1993).

METHODS

Broad Sampling Design Methods for assessing vegetation will be identical to those used for mapping during the previous two years. Maps, based on aerial photos taken during the Memorial Day Constant flows, will be attributed with vegetation layers (trees, shrubs, herbs, forbs, grasses and graminoids) and an assemblage type (sensu Brown et al. 1979). Methods for assessing

Kearsley and Ayers 6 return current channel morphology changes will involve repeat surveys of marsh transect lines for comparison with previous surveys.

Methodologies to be Utilized Mapping vegetation will be done in the same manner it has been done in 1995. We will be using the same 9 sites we have mapped in the previous two years (see table 1). Maps will be generated from 400% enlargements of the Memorial Day air photos. Vegetation polygons, based on similarity of appearance, will be drawn on mylar, with adequate copies made for 3 2-person crews.

Polygons will be attributed in the following manner: First, a complete species list will be generated for the polygon by walking through it and noting all species present, including current years' annuals. Second, a series of up to 5 circular plots, up to 6 meters in diameter, will be censused at randomly located points within the polygon. In these plots, percent foliar cover for all species in the plots will be noted, along with the diameter of the plot. Third, vertical structure will be measured by taking up to 8 point-contact measures at randomly located points within the polygon.

Polygons will be compared to data from 1995 using several approaches. First, a comparison of species lists will indicate the level of species turnover (number of species lost, number of colonizing species). No statistics will be needed for this type of analysis; either a species is present or it is not. Second, the percent foliar cover data will be used to compare changes in populations of obligate wetland species (as per Reed 1988) and other important riparian species such as Tamarix, Salix, Populus, and Baccharis. This data will be converted to a total foliar area measure (polygon size x percent cover) and compared, using polygons as the unit of study, to 1995 data with a paired t-test. Third, the percent foliar cover data will also be used to compare the progress of invasive exotics including Erianthus ravennae, Lepidium latifolium, Eragrostis curvula, and the annual bromes. These data will be analyzed in the same manner as wetland and important species. If data normality is a problem, both data types can be analyzed with a non-parametric equivalent. Finally, the vegetated area in each vegetation type, generated by TWINSPAN analysis, will be compared to the previous years data to test for changes in the relative cover of those types.

Survey data, from coordination with sandbar studies personnel, will compare the cross-sectional area of marsh transects in 1996 to those from the spring of 1995. A pair of vegetation personnel will accompany the surveyors to the sites where our studies overlap, lay out the marsh transects using tapes, chaining pins, and pinflags. They will then assist survey personnel by running rod for the transects and wherever else their help is needed. Elevation data from these surveys will be compared to previous surveys to assess whether the 20,000 cfs flows from the summer of 1995 have led to significant deposition or erosion in the return channels. We will calculate the areas between the transect profiles in 1995 and 1996 by calculating the areas between the two profiles.

Kearsley and Ayers 7 Logistics Support Requirements In order to carry out the survey part of the monitoring, we need only to have 2 people added to the survey trip scheduled for next spring. Data can be transferred from the NAU Sandbar database for the analyses we will need to do. The ground truthing trip, tentatively scheduled for August 1996 (for reasons relating to the timing of the delivery of the air photos from Horizons, Inc.) can be a GCES-run trip, as there are enough Botanist/Guides available who are willing to work for field assistant wages. I expect the trip will include 6 botanists for 16 days, based on the experiences from the just-completed mapping trip and last year's mapping trip. Tasks and Monitoring Timetable Activity*** Date

Initiation 1 October 1995 Begin work on integration of previous data into GCES-GIS 1 October 1995 Quarterly Report 1 April 1995 Spring tag-along with Sandbar Survey trip 14 April 1996 Quarterly Report 1 July 1996 Vegetation Mapping Trip 8 August 1996 Draft Final Report 15 September 1996 Final Administrative and Technical Report 1 October 1996

***Does not include 4 Transition Monitoring Workgroup meetings, which may be necessary.

DELIVERABLES As per the schedule above, the deliverables from this project will include quarterly, annual, and final reports as required. In addition, the final report will include attributed vegetation maps with comparisons to previous years' data and integrated data from fiscal years 1993 - 1995.

PERMITS Collecting permits will be required for voucher specimens for unknowns collected in the field. A research/study permit application is attached.

METADATA STANDARDS We will follow the metadata standards set forth in the request for proposals.

PRINCIPAL INVESTIGATORS Dr. Michael J.C. Kearsley will serve as Co-Principal Investigator on this project. He is fully qualified to serve as Co-P.I., having conducted GCES-II and Interim Flows studies, and published on the ecology of riparian plant species. Dr. Kearsley's responsibilities will include the coordination and orientation of field crews, coordination, packing and leading of field trips, performing and overseeing the entry, quality control, archiving, analysis, and interpretation of field data, and participation in the writing up of the results in reports and

Kearsley and Ayers 8 manuscripts for publication in professional journals. Dr. Kearsley will also be responsible for personnel issues related to the hiring and oversight of field crews.

Dr. Tina J. Ayers will serve as Co-Principal Investigator on this project. She is fully qualified to serve as Co-P.I., having conducted GCES-H and Interim Flows studies, and published extensively on plant . Dr. Ayers' responsibilities will include the identification and collection of plants in the field as unknowns. In addition, Dr Ayers will use her connections to others in her field to arrange for one or more other qualified systematists to accompany the trips. In the field, the trip taxonomists will identify (when possible) unknows plants, press specimens, and keep records of the collections.

LITERATURE CITED Brian, N.J. 1982. A preliminary study of the riparian coyote willow communities along the Colorado River in Grand Canyon National Park, AZ. No. Ariz. Univ. MS Thesis, Flagstaff.

Brian, N.J. 1988. Aerial photography comparison of 1983 high flow impacts to vegetation at eight Colorado River beaches. Glen Canyon Environmental Studies Rept. 20, Salt Lake City.

Brown, D.E., C.H. Lowe, and C.P. Pase. 1979. A digitized classification system for the biotic communities of North America with community (series) and association examples for the Southwest. Arizona-Nevada Academy Science 14 (Sup. 1): 1 - 16.

Campbell, C.J. and W. Green. 1968. Perpetual succession of stream-channel vegetation in a semiarid region. J. Ariz. Acad. Sci. 5: 86-98.

Carothers, S.W. and S.W. Aitchison. 1976. An ecological inventory of the Colorado River between Lees Ferry and the Grand Wash Cliffs. Grand Canyon National Park Colorado River Research Ser. 10, Grand Canyon.

Johnson, R.R., C.D. Ziebell, D.R. Patton, P.F. Ffolliott, and R.H. Hamre. 1985. Riparian ecosystems and their management: reconciling conflicting uses. Proc. First No. Amer. Riparian Conf., Tucson.

Kearsley, M.J.C. and T.J. Ayers. 1995. Continuing effects of Interim Flows on riparian plant assemblages in Grand Canyon National Park, Grand Canyon, AZ. Final technical and Administrative report. in prep

Koslowski, T.T., ed. 1984. Flooding and plant growth. Academic Press, Orlando.

Phillips, B.G., A.M. Phillips IH, M. Theroux, J. Downs, and G. Fryberger. 1977. Riparian vegetation of Grand Canyon National Park, Arizona. Mus. of Northern

Kearsley and Ayers 9 Ariz., Flagstaff. Unpublished map.

Phillips, B.G., R.A.Jolmson, A.M. Phillips III and N.J. Brian. 1986. Monitoring the effects of recreational use on Colorado River beaches in Grand Canyon National Park. Mus. Northern Ariz. Bull. Ser. 55, Flagstaff, AZ.

Pucherelli, M. 1988. Evaluation of riparian vegetation trends in the Grand Canyon using mulitemporal remote sensing techniques. U.S.D.I. Bureau of Reclamation Glen Canyon Environmental Studies Rept. No. 18. NTIS No. PB88-183488.

Reed, P.B. 1988. National list of plants that occur in wetlands alphabetized by common name and scientific name. Resource Management Group. Grand Haven, MI. Vol. 7. The Southwest.

Stevens, L.E. 1989. Mechanisms of riparian plant community organization and succession in the Grand Canyon, Arizona. Northern Arizona Univ. PhD Dissertation, Flagstaff.

Stevens, L.E. and G.L. Waring. 1988. Effects of post-dam flooding on riparian substrates, vegetation, and invertebrate populations in the Colorado River corridor in Grand Canyon, Arizona. U.S.D.I. Bureau of Reclamation Glen Canyon Environmental Studies Rept. No. 19. NTIS No. PB88183488/AS.

Stevens, L.E. and T.J. Ayers. 1992. The impacts of Glen Canyon Dam on riparian vegetation and soil stability in the Colorado River corridor, Grand Canyon, Arizona. Draft 1991 Annual Report to the Northern Arizona University NPS Cooperative Parks Studies Unit, Flagstaff. Unpublished.

Stevens, L.E. and T.J. Ayers. 1993. The impacts of Glen Canyon Dam on riparian vegetation and soil stability in the Colorado River Corridor, Grand Canyon, Arizona: 1992 Final Report. N.P.S. Cooperative Agreement CA 8000-8-0002.

Stevens, L.E., J.C. Schmidt, and T.J. Ayers. 1995 Geomorphic influences on fluvial marsh development along the dam-regulated Colorado River in the Grand Canyon, Arizona. Ecological Applications, in press

Turner, R.M. and M.M. Karpiscalc. 1980. Recent vegetation changes along the Colorado River between Glen Canyon Dam and Lake Mead, Arizona. U.S. Geol. Surv. Prof. Pap. 1132. Washington.

1Cearsley and Ayers 10 Budget Worksheet The following budget does not include the costs of logistics, which will be covered and coordinated by the Bureau of Reclamation GCES Office. Nor does it include technical survey support from the NAU sandbar studies group. Budget figured are presented here in thousands ("k") of dollars.

Item FY 1997 FY 1997 Total (est.) Personnel: Principal Investigators Kearsley $34.50k $35.12k ERE's (33%) 11.22k 11.56k Ayers (3/4 mo. @ $36,000 / 9 mo.) 3.00k 3.09k ERE's (33%) 1.02k 1.02k Field Technicians 3 Techs for 18 days @ 8 hrs./day @ $9.50 / hr. ERE's (8.5%) 4.11k 4.31k 0.35k 0.36k Total Personnel Costs $54.20k $55.46k $109.66k Equipment: Field equipment including notebook computer, mylar, pinflags, etc. $ 4.25k $ 1.25k $ 5.50k Office Supplies: Office supplies including storage media, software $ 1.25k $ 1.30k upgrades, color xeroxing, photofinishing and photo enlargements. Publication Page Charges (@ $75.00/pg) 1.50k 1.50k Total Supplies: $ 2.75k $ 2.80k $ 5.55k Travel: 1) To/from Flagstaff for meetings -0- -0- 2) For Monitoring Requirements -0- -0- 3) River Trips (1 veg trip plus 2 person tag-a-long on survey trip) -0- -0- 4) National Meetings for presentation of results Ecological Society of America (Aug. '95 in Providence, RI) $ 0.80k $ 0.80k Amer Soc. Plant Taxonomists (Aug. '95 in St. Louis, MO) 0.75k 0.75k

Total Travel $ 1.55k $ 1.55k $ 3.10k Analyses No analyses are required for this project -0- -0- -0- Subcontracts Contract with Deaver Herbarium for identifications and preparation $ 1.00k $ 1.00k $ 2.00k of specimens (20 hr. @ $50.00/hr)

TOTAL DIRECT COSTS _ $ 63.75k $ 62.06k $ 125.81k INDIRECT COSTS (20% OF DIRECT COSTS) _ $ 12.75k $ 12.41k $ 25.16k TOTAL COSTS $ 76.50k $ 74.47k $ 150.97k CURRICULUM VITA

Michael J.C. Kearsley

Permanent Mailing Address: 5557 White Horse Dr. Address: Dept. of Biol. Sci. Flagstaff, AZ 86004 Box 5640, N.A.U. (602) 773-9807 Flagstaff, AZ 86011 (602) 523-2381

Personal: Born in Boston, MA 14 September 1956 Married to Elizabeth Hall 15 June 1991

Education:

Northern Arizona University, Ph.D. Botany 1992. Major advisor: Thomas G. Whitham.

Northern Arizona University, M.Sc. Biology 1989. Major Advisor: Thomas G. Whitham.

Museum of Northern Arizona, Fern Mountain Field Ecology Program. 1982.

Dartmouth College, A.B. Biology Major. 1979.

Cornell University, Visiting Extramural Student in the Department of Entomology. 1978-1979.

Memberships:

Ecological Society of America American Society of Naturalists Entomological Society of America Society for the Study of Evolution

Professional and Relevant Experience:

1994-Pres Research Specialist / Adjunct Faculty. Department of Biological Sciences, Northern Arizona University, Flagstaff, AZ.

1991-1994 Research Specialist (Crew Leader / Data Analyst). Department of Biological Sciences / Glen Canyon Environmental Studies, Northern Arizona University. Flagstaff, AZ.

1990-1991. Research Assistant (Crew Leader / Data Analyst). Department of Geology / Glen Canyon Environmental Studies, Northern Arizona University. Flagstaff, AZ. Professional and Relevant Experience (Cont'd.)

1985-1990. Research Assistant. Department of Biological Sciences, N.S.F. Cottonwood Grants, Northern Arizona University. Flagstaff, AZ.

1984-1985. Teaching Assistant. Department of Biological Sciences, Northern Arizona University. Flagstaff, AZ.

1983-1984. Research Assistant. Department of Biology, Museum of Northern Arizona. Flagstaff, AZ.

1982-1983. Research Assistant. Department of Biological Sciences, Organized Research Office, Northern Arizona University. Flagstaff, AZ.

1980. Environmental Awareness Coordinator. Trail Camp Youth Conservation Corps, Coeur d'Alene National Forest. Coeur d'Alene, ID.

1979. Research Assistant. Department of Biological Science, N.S.F. Hubbard Brook Studies, Dartmouth College. Hanover, NH.

1977-1978. Research Technician. Cold Regions Research and Engineering Laboratories, U.S. Army Corps of Engineers. Hanover, NH.

1975. Biological Aide. Plumb Island Youth Conservation Corps, Plumb Island National Wildlife Refuge. Newburyport, MA.

Awards:

1991 Graduate Student Research Award. Sigma Xi Northern Arizona Chapter.

1985 Sigma Xi Grant-In-Aid of Research. "Developmental Resistance in Narrowleaf Cottonwood."

1983 Sigma Xi Grant-In-Aid of Research. "Tree- and site-specific determinants of herbivore success in the Narrowleaf Cottonwood (Populus angustifolia) galling aphid (Pemphigus betae) system."

Grants Contributed to:

1983 National Science Foundation. "Ecological and evolutionary significance of within- and between-plant variation in resistance to parasite attack." BSR-8303051 ($139,000).

1986 National Science Foundation. "Variation in host suitability and mechanisms of resistance to parasites in cottonwood." BSR-8604983 ($241,000).

1989 National Science Foundation. "Hybrid plants and hybrid zones as sinks for herbivores." BSR-8906391 ($335,000).

1995 Bureau of Reclamation / GCES. "A proposal to continue monitoring the effects on interim flows in riparian vegetation in Grand Canyon National Park, Grand Canyon, Arizona. ' ($94,000)

Service Activities:

1988 Graduate Student Representative. Biology Faculty Search Committee.

1986-1987. Secretary. Biology Graduate Students Association.

Publications:

1989 Kearsley, M.J.C. and T.G. Whitham. "Developmental changes in resistance to herbivory: Implications for individuals and populations." ECOLOGY 70 (2): 422-434.

1992 Kearsley, M.J.C. and T.G. Whitham. "Guns and butter: A no-cost defense against predation for Chrysomela confluens." OECOLOGIA 92: 556-562.

1993 Floate, K.D., M.J.C. Kearsley, and T.G. Whitham. "Elevated herbivory in plant hybrid zones: Chrysomela confluens, Populus and phenological sinks. ECOLOGY 74(7): 2056-2065.

1995 Kearsley, M.J.C. and T.J. Ayers. "Continuing effects of interim flows on riparian plant assemblages in Grand Canyon. Final Report (in prep.)

Invited Presentations:

1988 "Ontogenetic Resistance: Is It Real?" Workshop moderator, Western Forest Work Conference. Flagstaff, AZ.

Papers Presented:

1995 "Continuing effects of an altered flow regime on fluvial marshes in Grand Canyon." Ecological Society of America National Meeting. Snowbird, UT.

1994 "Effects of an altered flow regime on fluvial marshes in Grand Canyon National Park, Grand Canyon, AZ." Ecological Society of America National Meeting. Knoxville, TN

1993 "Effects of interactions among geomorphology, hydrology, and disturbance on riparian plant communities in Grand Canyon National Park." Ecological Society of America National Meeting. Madison, WI.

1993 "Effects of altered dam-regulated flow regime on riparian plant communities in Grand Canyon National Park." Arizona Riparian Council Annual Meeting. Rio Rico, AZ.

1989 "Effects of cottonwood host genotype and growth vigor on the performance of Chrysomela confluens. National Meeting, Entomological Society of America. San Antonio, TX.

1989 "Relative impacts of host plant vigor and genotype on the performance of a leaf skeletonizing beetle." Annual Meeting, Southwest Association of Biologists. Abiquiu, NM.

1988 "Narrowleaf cottonwood ontogeny and resistance to herbivory." Annual Meeting, Southwest Association of Biologists. Portal, AZ.

1988 "Narrowleaf cottonwood ontogeny creates a developmentally based mosaic of resisance to herbivory by galling aphids." National Meeting, Ecological Society of America. Davis, CA Papers Presented (Cont'd.)

1987 "Narrowleaf cottonwood ontogeny creates a developmentally based mosaic of resistance to herbivory by galling aphids." National Meeting, Entomological Society of America. Boston, MA.

1987 "Guns and butter: A no-cost self defense for Chrysomela confluens." Annual Meeting, Southwest Association of Biologists. Payson, AZ.

1986 "Is there a cost to self defense in Chrysomelid beetles?" National Meeting, Entomological Society of America. Reno, NV.

1985 "Developmental resistance in Narrowleaf Cottonwood." National Meeting, Entomological Society of America. Hollywood, FL.

1985 "Developmental resistance in Narrowleaf Cottonwood." Annual Meeting, Guild of Rocky Mountain Population Biologists. Ogden, UT.

1984 "Contrasting effects of age-related resistance on two herbivores of Narrowleaf Cottonwood. National Meeting, Entomological Society of America. San Antonio, TX.

1983 "Age-specific attack of Pemphigus betae on Narrowleaf Cottonwood (Populus angustifolia). Annual Meeting, Guild of Rocky Mountain Population Biologists. Crested Butte, CO. CURRICULUM VITA

Tina Jeanette Ayers

ADDRESS: Department of Biological Sciences, P. 0. Box 5640, Northern Arizona University, Flagstaff, AZ 86011; phone (602) 523-9482.

BIRTHDATE: 26 December 1957; Alexandria, Louisiana

EDUCATION: Ph.D. (Botany) - Systematics, 1986; University of Texas, Austin. B.A. (Botany) - General Botany, 1979; University of Texas, Austin.

MARITAL STATUS: Married; to Randall W. Scott

EMPLOYMENT AND TEACHING EXPERIENCE:

Assistant Professor & 1990-present Biological Sciences, NAU Curator, Deaver Herbarium

Assistant Research 1989-1990 Santa Barbara Botanic Garden, Scientist Flora of Santa Cruz Island Project --funded by The Nature Conservancy.

Research Assistant 1988-1989 Bailey Hortorium, Cornell University, Systematic Study of Palms, Anatomical and Chloroplast DNA research--funded by NSF.

Postdoctoral Associate 1987-1988 Bailey Hortorium, Cornell University, Cultivated Flora Project.

Postdoctoral Intern 1986-1987 New York Botanical Garden Bronx, New York

Assistant Instructor 1985-1986 University of Texas: Biology Department

Teaching Assistant 1982-1985 University of Texas: Biology/Botany

Staff Botanist 1979-1981 Camp, Dresser & McKee Environmental Sciences Austin/Denver PROFESSIONAL SOCIETIES: American Society of Plant Taxonomists International Association of Plant Taxonomists Willi Hennig Society The California Botanical Society Arizona Native Plants Society AWARDS:

1985 Harold Bold Teaching Award

GRANT HISTORY: Jun 1991 Impacts of Glen Canyon Dam on Riparian Vegetation. National Park Service. $101,820 (F) Jun 1992 Monitoring Effects of Interim Flows from Glen Canyon Dam on Riparian Vegetation. National Park Service. $262,410 (F) Jun 1992 Glen Canyon Dam Interim Flow Monitoring Program from Dam to Lee's Ferry. National Park Service. $4000 (F) Sep1992 Floristic Studies in Glen Canyon National Recreation Area. National Park Service. $2500 (F) Jul 1993 Systematics of Lysipomia. NSF, Research Planning Grant. 17,796 (F) Aug1993 Glen Canyon Dam Interim Flow Monitoring Program from Dam to Lee's Ferry, year 2. National Park Service. $4000 (F) Jan 1994 A Computerized Tutorial for Undergraduate Teaching of the Flora of Arizona. USDA. $80,000 (N) Jan 1994 NSF-REU extention on Systematics of Lysipomia. NSF. $5000 (F) Sep 1994 Monitoring Effects of Interim Flows from Glen Canyon Dam on Riparian Vegetation in Grand Canyon National Park. National Park Service. $94,000 (F) Jun 1995 Phylogenetics of Lysipomia: evolution in a Paramo endemic. NSF. $120,000 (P) Jun 1995 The Consortium of Herbaria in Arizona and New Mexico (CHAZNM): Development of a Computerized, Network-Accessible, Inter-Institutional, Herbarium Specimen Database. $69,163 (P). NAU ORG'91-92 Chloroplast DNA Evolution and Phylogeny of the Red-flowered Species of Lobelia (travel and supplies funded, capital equipment was frozen) NAU ORG'92-93 Floral Development and Pollen Morphology as Indicators of Common Ancestry in the Lobeliaceae (fully funded) NAU ORG'93-94 Phylogenetic Study of Lysipomia I: Speciation above and below the Huancabamba Depression (fully funded except travel 50% funded) NAU ORG'94-95 Phylogenetic Study of Lysipomia II: Resolution of conflicting hypotheses (fully funded) NAU OID'94-94 Macrophotography of Plant Material for Bio 415, Plant Taxonomy (fully funded) NAU Computer Services '95-96 A Multimedia Database for Teaching the flora and Ecosystems of the Southwestern United States, submitted with E. Taylor and T. Kolb, Forestry (fully funded) NAU ORG'95-96 Population Genetics of Circium wheeleri. (fully funded)

INVITED SEMINARS:

May 1989--University of California, Biological Sciences Department, Santa Barbara, CA. Nov 1989--Rancho Santa Anna Botanic Garden, Pomona, CA. Jan 1990--Northern Arizona University, Biological Sciences, Flagstaff, CA. Feb 1991--Northern Arizona University, Biological Sciences, Ecolunch. Dec 1991--Arizona State University, Botany Department, Tempe, AZ. Jan 1992--Arizona Native Plant Society, Flagstaff Chapter. April 1994-- Northern Arizona University, Biological Sciences, Ecolunch. May 1994--Uses of Arizona Natives--Sedona Native Plants Symposium (sponsored by Keep Sedona Beautiful) Nov 1994—Arizona Native Plant Society, Flagstaff Chapter. March 1995--Bureau of Land Management, Interagency Non-indigenous Plant Symposium, St. George, UT.

2 PROFESSIONAL CO1VIIVIITTEES:

1986-87 Associate Editor, Brittonia 1991-94 Publicity Committee, American Society of Plant Taxonomists 1994-96 Associate Editor, BioScience 1994-96 Treasurer, Arizona Native Plant Society, Northern Arizona Chapter

Reviewer: American Journal of Botany, Baileya, Brittonia, Sida, Systematic Botany, Journal of the Arizona- Nevada Academy of Science, NSF -- Division of Environmental Biology

PUBLISHED ABSTRACTS:

Ayers, T. J. 1986. The systematic implications of nectar spurs in the (). Amer. J. Bot. 73: 749. Ayers, T. J. 1988. The evolution of nectar spurs in Mexican lobeliads. Amer. J. Bot. 75: 158. Ayers, T. J. 1988. Reproductive biology in two spurred and two non-spurred Mexican lobeliads. Amer. J. Bot. 75: 58. Uhl, N. W. & T. J. Ayers. 1989. Carpoxylon macrospermum (Palmae, Incertae Sedis) and its relationships. Amer. J. Bot. 76: 741. Ayers, T. J. 1991. Preleminary phylogenetic and biogeographical studies of Lysipomia (Campanulaceae). Amer. J. Bot: 78: Bechtel, D. A., L. E. Stevens, M. J. Kearsley and T. J. Ayers. 1993. Geomorphic and hydrologic controls on riparian vegetation in the Grand Canyon, Arizona. Bull. Ecol. Soc. Am. Supp1.74:159. Ayers, T.J. 1994. Floral resupination in the Lobeliaceae: a twist on a twist. Amer. J. Bot. 81:140. Kearsley, M., L. Stevens, and T. J. Ayers. 1994. The effects of interim flows on fluvial marsh plant assemblages in Grand Canyon, AZ. Bull. Ecol. Soc. Am. Suppl. 74: A. Hindman and T. J. Ayers. 1995. Contractile roots as adaptations to life above 4000 meters. AZ-NV Acad. Sci. Suppl. 30: 21. L. E. Stevens and T. J. Ayers. 1995. Flooding and wetland development along a dam-regulated river. Bull. Ecol. Soc. Am. Suppl. 75: 254.

PUBLICATIONS:

Ayers, T. J. 1986. Systematics of Heterotoma (Campanulaceae: Lobelioideae). Ph.D. Thesis, University of Texas, Austin. Ayers, T. J. 1987. Three species from western Mexico new to Lobelia. Brittonia 39: 417-422. Ayers, T. J. 1988. Two Lobelia gypsophiles from Nuevo Leon, Mexico. Sida 13: 141-148. Ayers, T. J. & D. Boufford. 1988. Index to the types collected by H. H. Smith near Santa Marta, Colombia. Brittonia 40: 400-438. Ayers, T. J., R. W. Scott and B. L. Turner. 1989. Systematics of the two monotypic genera Mexianthus and Neohintonia (Asteraceae: Eupatorieae). Sida 13: 335-344. Ayers, T. J. 1990. Systematics of Heterotoma Zucc., and the evolution of nectar spurs in the New World Lobelioideae. Syst. Bot. 38:296-327. Ayers, T. J. 1991. The cultivated species of Asyneuma and Phyteuma. Baileya 24: 12-19. Ayers, T. J. 1993. Downingia and Lobelia. In: J. Hickman, ed., The Jepson Manual, Univ. of California Presss. Berkeley.

3 PUBLICATIONS (continued):

Stevens, L.E. and T. J. Ayers. 1993. The impacts of Glen Canyon Dam on riparian vegetation and soil stability in the Colorado River corridor, Grand Canyon, Arizona: final report. Submitted to NPS CPSU, Northern Arizona University, Flagstaff. Stevens, L.E. and T. J. Ayers. 1994. The effects of interim flows from Glen Canyon Dam on riparian vegetation in the Colorado River corridor downstream from Glen Canyon Dam, Arizona: draft final report. Submitted to NPS CPSU, Northern Arizona University, Flagstaff. Ayers, T., and L. Stevens. 1994. New and Noteworthy Reports: Arizona. Madrofio. 41: 228-229. Ayers, T., R. Scott, L. Stevens, A. Phillips, and K. Warren. 1995. Additions to the flora of Grand Canyon National Park. AZ-NV Acad. Sci. 28: 70-75. Junak, S., T. Ayers, R. Scott, and D. Young. 1995. A Flora of Santa Cruz Island. Santa Barbara Botanic Garden, Santa Barbara, CA. in press. Stevens, L.E., J. C. Schmidt, T. J. Ayers, B.T. Brown. 1995. Flow regulation, geomorphology, and Colorado River marsh development in the Grand Canyon, Arizona. Ecological Applications, in press. Ayers, T. In review (a). New species of Lysipomia from the paramo of southern Ecuador. Brittonia, submitted. Ayers, T. and A. Hindman. In review (b). Systematic significance of pollen morphology in the Andean endemic Lysipomia. Grana, submitted. Johnston, M. C. & T. J. Ayers. In review (c). Lobelia. In: M. Johnston and J. Henrickson eds., The Chihuahuan Desert Flora, submitted.

MANUSCRIPTS IN PREPARATION:

Ayers, T. J. and T. Lammers. Reevaluation of the Howellinae (Lobeliaceae), to be submitted to Brittonia. K. Huisinga and T. Ayers. Plantaginaceae for Vascular Plants of Arizona Project, to be submitted to Journal of the AZ-NV Academy of Science. Ayers, T. J. and A. Hindman. Evolution of contractile roots in Lysipomia, a paramo endemic, to be submitted to American Journal of Botany. Ayers, T. J. Systematic significance of leaf hydathodes in the Lobeliaceae, to be submitted to Plant Systematics and Evolution. Ayers, T.J. Monograph of Lysipomia (Lobeliaceae), to be submitted to Systematic Botany Monographs. Ayers, T.J. Origin, evolution, and biogeography of Lysipomia, a genus endemic to Andean paramo and puna, to be submitted to Cladistics. Ayers, T. J. A phytogeographic re-evaluation of the Lobelia berlandieri A. DC. species complex in Northern Mexico, to be submitted to Southwestern Naturalist.

4

APPENDIX A

• GRAND CANYON NATIONAL PARK GLEN CANYON NATIONAL RECREATION AREA

RESEARCH/STUDY PERMIT APPLICATION

Date 20 Aug. 1995 Project start date: j Ort. 1995

PROJECT TITLE

Transition monitoring for riparian vegetation in thp Colorado River corridor of Grand Canyon National Park

PRINCIPAL INVESTIGATOR

Name: Michael J.C. Kearsley / Tina J. Ayers Northern Arizona University Affiliation:

Address: P.O. Box 5640. Northern Arizona UniVprgitv

Flagstaff, AZ 86011

Telephone # (520) 523-2381 e-mail # [email protected]

COLLECTIONS

Type and number of specimens to be taken: Voucher specimens for unknown species 1 from study plots.

DESCRIPTION OF PROJECT: A complete proposal is required for all studies conducted in Grand Canyon NP and Glen Canyon NRA. Attach a proposal for the proposed study• following guidelines set forth in Appendix B.