At Clarke & Pine, Hoosier Prairie, Indiana Dunes State

Home , Cypripedium candidum, Epigaea repens

LONG-TERM CHANGES IN POPULATIONS OF ENDANGERED AND THREATENED PLANTS AT CLARKE & PINE, HOOSIER PRAIRIE, INDIANA DUNES STATE PARK, AND TOLLESTON RIDGES/GIBSON WOODS NATURE PRESERVES

2005

Marlin Bowles, The Morton Arboretum Michael Jones, Christopher Burke Engineering Ltd. Jenny McBride, The Morton Arboretum

SUMMARY

Goals and methods - To assess their current status and long-term changes, we re-sampled 65 populations of 60 Indiana endangered or threatened plant species at the Clarke & Pine, Hoosier Prairie, Indiana Dunes, and Tolleston Ridges/Gibson Woods Nature Preserves. These populations were initially sampled between 1986 and 1989, which allowed assessment of their long-term changes. To help understand factors that might be contributing to these changes, we also re-surveyed and analyzed structural changes in associated plant community vegetation at Clarke & Pine and Tolleston Ridges.

Status and change- We were able to assess the status of about 78 % of the original populations. Another 8 % were present in low numbers and were not quantified, and the status of 14 % was unknown. Among all plant populations studied, 35 % appeared to be stable over time, while 37 % declined and 6 % increased either in numbers or other critical measures, such as % flowering or genet size. These results suggest that most populations either have been stable or have declined, and that populations are more likely to decline that to increase in numbers over time. At Clarke & Pine, 4 of 18 species underwent significant declines, while 7 other species were not present or too rare to quantify. Among these species, Hypericum kalmianum had the greatest decline. Paper birch, as well as the alien buckthorn Rhamnus frangula also increased significantly. At Hoosier Prairie, 4 species, including Eriophorum angustifolium, could not be relocated and their status is unknown. Several other species were either too rare to quantify or their populations have shifted. The orchid Malaxis unifolia increased significantly. At Indiana Dunes State Park, dynamic changes appear to be occurring in many shoreline populations, while others are either recovering from deer browsing, or may have been extirpated. Polygala paucifolia appears to be extremely vulnerable to human impacts. At Tolleston Ridges/Gibson Woods, the orchids Cypripedium reginae and C. parviflorum declined, while the sedges Carex aurea and C. richardsonii could not be relocated.

Causes of change - Lack of experimental treatments limits understanding causes of change in many populations, and multiple factors may have contributed to changes at different sites. Low frequency of fire appears to be a critical factor in change among some species at Clarke & Pine, including an increase in woody vegetation. Ongoing fire management at Hoosier Prairie also may be driving successional changes in vegetation and shifting spatial patterns among species, including both perennials and short lived species that respond rapidly to fire. At Indiana Dunes, population sizes of shoreline-adapted species may fluctuate in response to lake level cycles that drive sand erosion and deposition, as well as vegetation succession, while human impacts and over browsing from deer appear to have impacted other species. Lack of prescribed burning has lead to decline of species and structural shifts of vegetation at Tolleston Ridges and Gibson Woods. Periodic natural fluctuations in population sizes, as well as natural long-term changes in vegetation structure may also be important causes of changes that were detected in this study.

Recommendations - More frequent monitoring and greater replication of plots (when possible) are needed to better understand how rare plant populations are changing at these study sites. Sampling frequency should be based upon the longevity and life history strategy of different plants, with more frequent monitoring for short-lived species. To better understand population trajectories, annual demographic monitoring would be required, which is being used for Cirsium pitcheri. Monitoring within an experimental framework, such comparing between burned and unburned treatments, also may be needed to better understand how to maintain some populations. Increased use of fire appears to be a critical need for recovery of species at Tolleston Ridges and Gibson Woods, and it may be important in long term maintenance of vegetation at Clarke & Pine.

Acknowledgments - We thank Dan McDowell for his assistance with most of the field work associated with this report, and for sharing his knowledge of rare plant populations in the Indiana Dunes. We also thank Sandy Obrien for her assistance, Tom Post for management information and John Bacone for coordinating the project. This work was made possible by a grant from the National Oceanic and Atmospheric Administration and the Indiana Department of Natural Resources, Lake Michigan Coastal Program.

Table of Contents Page INTRODUCTION ...... 1

METHODS ...... 1

RESULTS AND DISCUSSION ...... 2 Clarke & Pine ...... 2 Hoosier Prairie ...... 3 Indiana Dunes ...... 4 Tolleston Ridges/Gibson Woods ...... 5

CONCLUSIONS ...... 6

REFERENCES ...... 8

Figures 1) Twenty-year fire frequencies at Clarke & Pine. 2) Lake Michigan lake levels 1960-2005.

Appendices I. Original maps with species plots and baseline locations for Clarke & Pine, Tolleston Ridges (Shell), and Hoosier Prairie II. Clarke & Pine Species and vegetation changes III. Hoosier Prairie species changes. IV. Indiana Dunes species changes. V. Tolleston Ridges/Gibson Woods species and vegetation changes. VI. Aerial and topographic shape files, meta data, and GPS Coordinates

INTRODUCTION

The Indiana Dunes region of northwest Indiana has one of the Midwest’s greatest concentrations of rare plant species (Peattie 1930, Pepoon 1927, Deam 1940, Wilhelm 1990 Swink & Wilhem 1994), as well as remnant natural areas (Bacone 1979, Kurz 1979, Bowles 1989). This wealth of biological diversity probably survived due to the extreme habitat conditions of its sand soils, which prevented wide-scale development or conversion to agriculture (Moore 1959, Bowles et al. 1982, Homoya et al 1985).

To provide a basis for long term monitoring of rare plant species in this area, permanent sampling plots were established for Federal and State of Indiana endangered and threatened plants on Federally owned units of the Indiana Dunes National Lakeshore in 1984 and 1985 (Bowles et al. 1985, 1986a, 1986b). As a continuation of this project in 1986, 66 populations of 60 Indiana endangered or threatened plant species were studied at the Indiana Dunes State Park, as well as at the satellite areas Clarke & Pine, Hoosier Prairie, and Tolleston Ridges/Gibson Woods Nature Preserves (Bowles 1987). Although some of these species have been re-monitored (Bowles 1988, 1989, 1990), there has been no comprehensive survey of the status of all species that were originally sampled. In 2004, we investigated the status of species at Clarke & Pine, Indiana Dunes, Hoosier Prairie, and Tolleston Ridges/Gibson Woods Nature Preserves by attempting to relocate and re-sample their original plots. To better understand detected changes, we also re-sampled changes in vegetation structure at Clarke & Pine and at Tolleston Ridges-Gibson Woods, and also assessed site burn frequencies for these sites. Although disturbance processes are important for many of the shoreline species (Bowles et al1990), we expected that rare species in prairie and savanna habitats might be more stable temporally if vegetation structure is also stabilized by fire management.

METHODS

Original data were based on random or arbitrary population samples that were permanently marked, surveyed and mapped. We used this location information to re-sample plots for data that could be to compared statistically with original data sets. We also collected GPS coordinates for all plots that were relocated, as well as for some new populations that were sampled. The coordinates are provided with GPS shape files for these species.

At Clarke & Pine, most plots were associated with baselines established on the main north and south ridges. Appendix I contains original plot and baseline maps. The 500 m south ridge baseline also served as a transect for groundlayer vegetation plots, which were also resampled (N = 50 1-m2 plots). Vegetation canopy cover was also monitored in association with Pinus

banksiana on a ridge located south of the south baseline. At Tolleston Ridges, Cypripedium calceolus var. parviflorum and Carex richardsonii, as well as dry-mesic prairie groundlayer plots (N = 10 1/4-m2 plots), were re-sampled along the central ridge. Demographic plots for Cypripedium reginae and canopy light values were sampled on the south ridge at Tolleston. Groundlayer vegetation plots (n = 28) were also resampled along seven North-South transects from a baseline that bordered the south ridge at Tolleston Ridges. Although no surveyed baseline was established Indiana Dunes, a 2000 m length of Trail #2 was used to monitor Botrychium species, Milleum effusum and Poa alsodes. We also compiled and mapped the coverage of 1984-2005 fires at Clarke & Pine (Figure 1), and accessed historic Lake Michigan lake levels from (Figure 2).

Chi-square analysis was used to test for changes in species abundance among different plot categories over time. T-tests were used to compare mean differences in abundance between original and 2004 data, while one-way ANOVA was used to test differences among data sets that included more than two time frames.

RESULTS AND DISCUSSION

Clarke & Pine

Of the 18 species originally sampled at Clarke & Pine, seven appeared to be stable, while four underwent significant declines in at least on plot (Appendix II) Arenaria stricta declined from 75 % to 10 % plot frequency on the north baseline, but it increased from about 25 % to 40 % frequency on the south baseline. Carex richardsonii also declined from greater than 9 to fewer than 1 flowering ramets/m2 on the north baseline. Both of these declines may have resulted from a late spring burn in 2004 and therefore could represent temporary effects that might not be important for long-term persistence of these perennial species. However, Hypericum kalmianun dropped from 19 % to 2 % frequency along the south baseline and appears to have essentially disappeared from the site. Rhus aromatica var. arenaria also declined from about 34 % to 20 % plot frequency but still remains a common plant. Although Buchnera americana did not change significantly, > 50 % of its stems were browsed by eastern white-tailed deer in 2004. Browsing was not noted on this species in 1986.

Seven other species that were monitored in 1986 either could not be found, or occurred in such low numbers that their population changes could not be quantified. Carex eburnea originally occurred in association with Pinus banksiana, and may have disappeared as the stand matured (see below). Eleocharis geniculata and Eleocharis pauciflora appear in late summer as their habitat is exposed by drawdown of the water table in swales. Although Eleocharis pauciflora

was observed, the high water table in late 2004 prevented quantitative sampling. The status of Eleocharis geniculata is unknown; however it may remain extant. Orobanche fasciculata could not be located, but it is usually rare and might easily have been overlooked. Platanthera hyperborea originally occurred under shade of native shrubs, as well as the alien Rhamnus frangula, and apparently disappeared after this woody vegetation was removed by management. Utricularia purpurea was observed in the same vicinity in which it was originally sampled, but only a few flowering plants were present and the status of nonflowering individuals is unknown. Carex alata could not be relocated and we are unaware of its current status.

Only one study species, Betula papyrifera, underwent a significant increase. This tree increased in abundance from about 5 to 25 trees/ha, while its stem numbers (each tree has multiple stems) increased by an order of magnitude across all size classes. For example, stems in the 1-3 cm size classes increased from fewer than 25 stems to more than 200 stems/ha. There was no change in abundance of smaller stems, which had less than 15 % frequency in groundlayer plots in 1986 and inn 2004. This suggests that B. papyrifera has not increased in abundance as a component of groundlayer vegetation. Although there was no change in stem numbers of Pinus banksiana, there was a substantial shift in the structure of its population. The number of stems less than 0.5 dm in diameter dropped 40 %, while stems greater than 1 dm increased by 30 %, indicating an increase in the size of trees in this maturing population. Vegetation structure in the stand also changed, with a 30 % increase in savanna tree cover, a 30 % decline in cover of prairie and a 60 % increase in cover by Rhamnus frangula. On the south baseline, Rhamnus frangula also increased significantly from less than 25 % to greater than 45 % plot frequency, indicating increasing abundance as a component of groundlayer vegetation. Spiranthes magnicamporum was not originally monitored but noted as present. In 2004 we observed this orchid throughout the preserve; but it is unknown whether the size of the population has changed over time.

The substantial increase in paper birch and buckthorn at Clarke & Pine probably resulted from the low frequency (~ 2 fires in 20 years) at which prescribed burns occurred at this site. However, we found no significant declines in the number of species present in groundlayer plots, a key indicator of infrequent burning of prairie vegetation (Bowles & Jones 2004). This suggests that if the increases in paper birch and buckthorn are a response to low fire frequency, impacts on groundlayer herbaceous vegetation are not yet detectable. However, the increase in abundance of buckthorn appears to be a critical threat to the preserve.

Hoosier Prairie

Five of the 12 species originally monitored at Hoosier Prairie appear to have remained stable

over time (Appendix III). One species, the cottongrass Eriophorum angustifolium appears to have declined as it could not be relocated within its former habitat. However, it was observed elsewhere in the preserve in 2004 (T. Post, pers. comm.). The status of two additional sedges, Carex conoidea and Rhynchospora globularis var. recognita is unknown. They could not be relocated in plots on the NIPSCO right-of-way, where they may have disappeared with vegetation succession or after management of the ROW. The status of Polygonum caryei is also unknown, as it was not observed in former habitat. Corydalis sempervirens and Geranium bicknellii were present in original habitat, but there were too few plants to quantify their abundance. These short-lived species may appear in larger numbers in response to fire.

Several species may be undergoing spatial shifts and changes in density. Polytaenia nuttallii occupied a larger area than indicated by the 1986 sampling plot, but fewer plants were present, and the % of flowering plants was lower than in 1986. Satureja arkansana no longer occurred in the original sample plot, but occupied a larger adjacent area at lower densities in 2004. The orchids Cypripedium candidum and Platanterha flava var. herbiola still occur within original sample plots. Both appear to have declined moderately since the late 1980's, but they have not dropped below original sample numbers. One orchid species, Malixis unifolia, appears to have increased significantly in population size and numbers. This species now occupies a much larger habitat area that originally known, and increased in plot density as well as from about 20 plants to probably over 100 plants in the population.

Indiana Dunes

Of the 29 species populations monitored in the Indiana Dunes nature preserve, 11 appear to be stable, while 15 have declined, one has increased and the status of two is unknown (Appendix IV). The single species for which we recorded a substantial increase, Poa alsodes, occurs in mesic forest habitat along Trail #2. The inflorescences of this early flowering bluegrass disarticulate soon after flowering, which may have caused observers to record fewer plants in 1986. Milleum effusum, which co-occurs with Poa alsodes, is more easily detected and remained stable. Both species may benefit from increased light availability and disturbance along the trail edge. Two grape ferns, Botrychium matricariaefolium and Botrychium oneidense, also occur along Trail #2. Botrychium matricariaefolium declined from greater than 25 plants seen in 1984 and 1986 to 5 plants seen in 1990 and in 2004. However, it may be present in extensive potential habitat. Botrychium oneidense, in contrast, appears to have remained stable.

Six of the declining species occur in dynamic lakeshore habitats. In 2004, the annual Cakile edentula occurred at the lowest densities ever recorded for most plots. Euphorbia polygonifolia,

an annual that often occurs with Cakile edentula, was present in extremely low numbers that could not be quantified. In blowout habitat, Salix syrticola dropped from about 12 stems/m2 in 1986 and 1990 to less than 3 stems/m2 in 2004, while Potentilla anserina disappeared from an adjacent plot in 1989. In the two latter cases, sand deposition appears to have caused the changes, as the loss of Potentiall anserina was due to sand burial of its panne habitat (Bowles 1990), and we observed sand burial of Salix syrticola. Lake Michigan water levels since 1999 have been at their lowest since the 1960's (Figure 2), which may be increasing shoreline sand deposition. As a result, the observed population changes could be cyclic in response to fluctuating lake levels and sand movement, and may not signal threats to the status of these species unless they are exacerbated by human impacts (Bowles et al1990). Cirsium pitcheri declined from census numbers above 90 individuals in the late 1980's to fewer than 10 individuals. This poor competitor may be declining from ongoing plant succession in the plot area. However, adjacent habitat in the big blowout still supports hundreds of plants. Juniperus communis var. depressa occurs in more protected blowout habitat and appears to be more stable because of its longevity. This species underwent a significant shift in population structure due to past over-browsing by deer, followed by recovery. The population sample now contains a much greater proportion of juveniles than in 1986, and plants are slightly smaller in size. We were unable to relocate the grass Oryzopsis asperifolia, which occurred on the steep wooded slope of a blowout. This population may have been completely lost due to dune movement.

Visitor use may be impacting some species on steep fragile slopes. In north facing bluff habitat along Dune Creek, Melampyrum lineare declined from about 190 to 40 plants, but its ramet densities remained unchanged. In flatwoods habitat, it declined from almost 250 plants to about 25 plants, and also dropped significantly in plant density. Visitor use could be affecting this species by causing erosion of its sand bluff habitat along Dune Creek, but we saw no evidence of similar impacts in flatwoods. Erosion from foot traffic may be affecting Polygala paucifolia, or at least poses a threat to this species. The white population occurs on the steep slope of Mt. Jackson, and appears to have declined in total ramet densities, but not the density or percentage of flowering stems, which actually increased. The pink population, which occurs on less vulnerable habitat, did not decline in overall density and increased in flowering density and the percentage of flowering stems. Pyrola rotundifolia declined about 50% in plot frequency in habitat at Dune Creek, and also could be affected by erosion from foot traffic.

The status of three additional species is unclear. Chimaphilla umbella could not be reclocated, and may have disappeared due to past impacts from eastern white-tailed deer. Epigaea repens also could not be relocated in former plots, but it was present in nearby habitats - probably due to the rhizomatous growth habit of this species. Platanthera flava var. herbiola could not be relocated, but its habitat was covered by a tree fall in the late 1980's and it may be present

elsewhere in extensive flatwoods habitat (Bowles et al. 1990). Tolleston Ridges/Gibson Woods

Only four species were originally monitored in these preserves and all declined (Appendix V). The sedges Carex aurea and Carex richardsonii, as well as their associated prairie vegetation, could not be relocated. Both appear to have disappeared due to fire protection, as there was no evidence of recent prescribed burns in their habitats. In dry-mesic prairie habitat that formerly contained Carex richardsonii at Tolleston, the relative abundance of graminoid vegetation dropped by 50 %, while woody vegetation, including Rhamnus frangula, increased by over 300 % - thereby causing the structural deterioration of this vegetation.

The lady’s slipper orchids Cypripedium calceolus var. parviflorum and Cypripedium reginae also declined at Tolleston Ridges due to habitat alteration from lack of prescribed burns. In 1986, Cypripedium calceolus var. parviflorum was monitored in a 1/10 ha area, where 40 plants were located, with over 120 ramets. In 2004, only four plants were present. This represents a decline from 400 to fewer than 50 plants/ha, and from greater than 1200 ramets/ha to fewer than 200 ramets/ha. The Cypripedium reginae population at Tolleston Ridges averaged about 130 plants between 1992 and 1999, but dropped to fewer than 100 plants in 2004. This was accompanied by a significant decline from 40 % to less than 20 % canopy openness, which appears to have been caused by lack of burning. Both the number of flowers per ramet and % fruit set per plant are positively correlated with canopy openness, indicating that the population can be expected to continue to decline without fire management.

After burning and brush clearing in early 2005, canopy light levels in mesic savanna at Tolleston were increased to greater than 25 %, and a second Cypripedium reginae population census found 121 plants. This indicates that most plants had persisted despite the increasing brush cover. However, groundlayer sampling in 2005 indicated that significant shifts occurred in vegetation structure and species composition. Although plot richness of native forbs increased over time, this was primarily due to an increase in abundance of shade-adapted species. In contrast, most prairie species, especially grasses, declined in plot frequencies. Accompanying changes included a significant increase in plot richness of woody vegetation species and a decline in plot richness of graminoid species.

CONCLUSIONS AND RECOMMENDATIONS

Our re-sampling of endangered and threatened plants in five Indiana nature preserves quantified the status of about 78 % of the original populations. Another 8 % were present in low numbers and were not quantified, and the status of 14 % was unknown. Among all plant

populations studied, 35 % appeared to be stable over time, while 37 % declined and 6 % increased either in numbers or other critical measures, such as % flowering or genet size. These results suggest that most populations either have been stable or have declined, and that populations have been more likely to decline than to increase.

Lack of experimental treatments limits understanding causes of change in many populations, and multiple factors may have contributed to changes at different sites. Vegetation succession and shoreline sand erosion and deposition, as well as human impacts, no doubt strongly affect long-term changes in populations of plants adapted to shoreline habitats (Bowles et al 1990). Because these processes may be driven by lake level cycles that operate over decades, long term monitoring is needed to understand how disturbance-adapted species respond to these cycles, and whether human impacts exacerbate the effects of cyclic extremes. More direct human impacts may be affecting species that occupy steep slopes on normally forested dunes. The most critical of these species is Polygala pauciflolia. This northern species also could be declining with increasing temperatures associated with global warming.

Past over-browsing by eastern white-tail deer may have contributed to loss of several plants from the Indiana Dunes. The most obvious result of this impact is the shift in population structure of Juniperus communis var. depressa caused by loss of larger adult plants. Chimaphilla umbellata may have declined from deer impacts but there is no direct evidence. We also observed browsing impacts on Bucknera americana at Clarke & Pine in 2004.

The long history of prescribed burning at Hoosier Prairie may be driving long-term changes and spatial shifts in plant populations, as well as appearances of short-lived species that respond positively to fire. Lack of prescribed burning appears to have lead to loss or serious decline of plant species populations and shifts in vegetation structure at Tolleston Ridges and Gibson Woods. The most direct evidence is the loss of associated prairie vegetation structure and increasing canopy cover in habitats for these species. Long-term increases in woody vegetation, including both the state listed paper birch and the alien Rhamnus frangula, have also occurred at Clarke & Pine, but there is little direct evidence for impacts at the species level. Nevertheless, these changes pose a long-term threat to the preserve and may be regulated by fire frequency. Fire appears to cause direct mortality of paper birch stems, but also stimulates re-sprouting, and it is unknown as to how it affects population structure of this species.

More frequent monitoring and greater replication of plots (when possible) are needed to better understand how rare plant populations are changing at these study sites. Sampling frequency should be based upon the longevity and life history strategy of different plants, with more frequent monitoring for short-lived species. To better understand population trajectories, annual

demographic monitoring would be required, which is being used for Cirsium pitcheri. Monitoring within an experimental framework, such as comparing between burned and unburned treatments, may be needed to better understand how to maintain some populations. Increased use of fire appears to be a critical need for recovery of species at Tolleston Ridges and Gibson Woods, and it may be important in long term maintenance of vegetation at Clarke & Pine.

REFERENCES

Bacone, J. 1979. Shell Oil Dune and Swale, a Report on a Natural Area. Indiana Department of Natural Resources, Division of Nature Preserves. Indianapolis.

Bowles, M. 1987. The status of Indiana Special Concern Vascular Plants at Indiana Dunes, Clarke & Pine, Shell, Gibson Woods, Hoosier Prairie, and Tefft Savanna Nature Preserves, Lake Co., IN. The Morton Arboretum, Lisle, IL.

Bowles, M. 1988. Special floristic elements at the Indiana Dunes National lakeshore: new species monitoring and update of selected existing populations. The Morton Arboretum, Lisle, IL.

Bowles, M. 1989. Status Report on Endangered and Threatened Plants of the Indiana Dunes National Lakeshore: new species monitoring and update of selected existing populations. The Morton Arboretum, Lisle, IL.

Bowles, M. 1990. Status Report on Endangered and Threatened Plants of the Indiana Dunes National Lakeshore: new species monitoring and update of selected existing populations. The Morton Arboretum, Lisle, IL.

Bowles, M. W. Hess, & M. DeMauro. 1985. An Asessment of the Monitoring Program for Special Florisitc Elements at the Indiana Dunes national Lakeshore: Phase I. The Endangered Species. The Morton Arboretum, Lisle, IL.

Bowles, M. W. Hess, & M. DeMauro. 1986a. An Asessment of the Monitoring Program for Special Florisitc Elements at the Indiana Dunes national Lakeshore: Phase I. The Threatened Species. The Morton Arboretum, Lisle, IL.

Bowles, M., W. Hess, M. DeMaruo & R. Hiebert. 1986b. Endangered plant inventory and monitoring strategies at the Indiana Dunes National Lakeshore. Natural Areas Journal 6(1):18- 26.

Bowles, M., J.K. Castle, & T. Post. 1982. Management Plans for Clarke & Pine and Shell- Gibson Nature Preserves, Lake Co., IN. The Natural Land Institute, Rockford, IL.

Bowles, M.L., M.M. Demauro, N Pavlovic, & R.D. Hiebert. 1990. Effects of anthropogenic disturbances on endangered and threatened plants at the Indiana Dunes National Lakeshore. Natural Areas Journal 10:187-200.

Bowles, M. & M. Jones. 2004. Long-term changes in Chicago region prairie vegetation relation to fire management. Chicago Wilderness Journal 2(2):7-16.

Deam, C.C. 1940. Flora of Indiana. Department of Conservation, Division of Forestry. Indianapolis.

Homoya, M.A., D.B. Abrell, H.R. Aldrich & T. Post. 1985. The natural regions of Indiana. Proceedings of the Indiana Academy of Science 94:245-268.

Kurz. D. 1979. An Inventory of Natural Areas in the Indiana Coastal Zone Study Area. Indiana State Planing Services Agency Technical Report No. 302. Indiana Department of Natural Resources, Indianapolis & Natural Land Institute, Rockford, IL.

Moore, P.A. 1959. The Calumet Region: Indiana’s Last Frontier. Indiana Historical Bureau. Indianapolis.

Peattie, D.C. 1930. Flora of the Indiana Dunes. Field Museum of Natural History. Chicago, IL.

Pepoon, H.S. 1927. Fora of the Chicago region. Chicago Academy of Science and Natural History Bulletin 8:1-576.

Swink, F. & G. Wilhelm. 1994. Plants of the Chicago region. Indiana Academy of Science. Indianapolis, IN.

Wilhelm. G. 1990. Special vegetation of the Indian Dunes National Lakeshore. Indiana Dunes National Lakeshore Research Program, Report 90-02. National park Service, Department of the Interior.

Figure 1. Fire frequencies from 1984-2005 for Clarke & Pine Nature Preserve

Figure 2. Lake Michigan Lake Levels 1960-2005. Source: http:www.Ire.usuace.army.mil/index.cfm.

Appendix I. Original site maps indicating locations of plant population plots or community transects.

Figure 1. Clarke & Pine Nature Preserve Figure 2. Tolleston Ridges (Shell) Nature Preserve Figure 4. Hoosier Prairie

Appendix II. Clarke & Pine Nature Preserve Species Location & Q.# Plot Data Change

Arenaria stricta North BL #1 20 m-sq plot % frequency/m-sq decline Arenaria stricta South BL #2 500 m-sq plot % frequency/m-sq stable Aster ptarmicoides South BL 500 m transect - 99 plots % frequency/m-sq stable Betula papyrifera South BL 500 m transect - 10 plots genet density per 1/40 ha increase South BL 500 m transect - 10 plots stem size class density per 1/10 ha increase South BL 500 m transect - 99 plots % frequency/m-sq stable Buchnera americana North BL #1,2,3 3 tranects (33-m) total stems, density & % browsed stable Carex aurea South BL #1 2 m x 11 m transect stem density/m-sq stable Carex eburnea South ridge #1 50-m transect census unknown Carex richardsonii North BL #1 1 m x 25 m transect culm/stem density/m-sq decline Eleocharis geneiculata South BL #1 1 m x 20 m transect frequency by dm-sq present?? Eleocharis pauciflora South BL #1 1 m x 20 m transect frequency by dm-sq present Gerardia skinneriana South BL # 50-m transect stem density/m-sq stable Hypericum kalmianum South BL 500 m transect - 99 plots % frequency/m-sq decline Orbanche fasciculata South BL #1,2 m-sq stem density/m-sq unknown Pinus banksiana South ridge #1 50-m transect total census by size class stable Platanthera hyperborea North BL #1,2 m-sq, 20 m transect ramet density decline Rhamnus frangula (alien) South BL 500 m transect - 99 plots % frequency/m-sq increase Rhus aromatica arenaria South BL 500 m transect - 99 plots % frequency/m-sq decline Spiranthes magnicamporum None None None Present Utricularia purpurea South BL #1 1 m x 17 m transect flowering ramet density/m-sq Present Carex alata #1,2 1 m-sq, 1 m by 5 m transect genet & culm density unknown Community transect South 500 m transect - 99 plots Species density/m-sq by group increase Community transect Pinus banksiana 5-m transect Structural cover stable

Arenaria stricta 80

70 1986 60 2004

50 2 40

30 Frequency/m 20

0 North (P <.001) South (P = 0.092) Baseline Location (X 2 Probability)

Arenaria stricta Clarke & Pine

North Baseline: 20 m2 plot (% frequency by m2)

South Baseline: 500 m2 plot (% frequency by m2, randomly subsampled in 2001)

Aster ptarmicoides 45 40 35 30 25 20

Frequency 15 10 5 0 2 1986 X P = 0.374 2004

Aster ptarmicoides Clarke & Pine

South Baseline: 500 m transect (% frequency in 99 1-m2 plots)

Betula papyrifera Tree (genet) Density Betula papyrifera G roundlayer Frequency 15 45

40 12 35

30 9 25

20 6 15 Trees (genets)/ha % frequency in groundlayer % frequency 10 3 5

0 0 1986 2004 Z-test P = 0.001 1986 X2 P = 1.00 2004

Betula papyrifera Stem Density in 1986 Betula papyrifera Stem density in 2004

25 350

300 20

250

15 200

150 10 Stems per ha 100 Stems per ha Stems

5 50

0 0 2 3 4 5 6 7 8 9 0 1 2 ------1 1 1 -2 -3 -4 -5 -6 -7 -8 -9 0 1 2 - - - 1 2 3 4 5 6 7 8 -1 -1 -1 >1 >2 >3 >4 >5 >6 >7 >8 0 1 9 0 1 >9 > > > > > > > > > 1 1 > > >1 >1

Size class in cm Size class in cm

Clarke & Pine Betula papyrifera South Baseline: 500 m transect (genet density in ten 1/40 ha plots, stem density by size class in ten 1/10 ha circular plots, groundlayer % frequency in 99 1m2 plots). Change in Buchnera americana 60

50 Ramets % browsed 40

20 Total stems & % browsed 10

0 1986 2004

4.5 Change in Buchnera americana 4

3.5

2 2.5

2 Stems/m 1.5

0.5

0 1986 t-test P = 0.780 2004

Clarke & Pine Buchnera Americana

North Baseline: three cumulative 33 m transects (Total and % browsed, stems/m2) Carex aurea 8

5 + s.e. 2 4

3 Plants/m

0 1986 2004 t-test P = 0.645

Clarke & Pine Carex aurea

South Baselin: 2 m x 11 m transect (ramet density/m2)

14 Carex richardsonii 12

6 Flowering ramets/m2 4

0 1986 2004 t-test P = 0.012

Clarke & Pine Carex richardsonii

North Baseline: 1 m x 25 m transect (Flowering ramet density/m2) 25 Gerardia skinneriana 20

15 2

5 Mean density/m

0 1986 t-test P = 0.064 2004

Clarke & Pine Gerardia skinneriana

South Baseline: 1986: 20 m transect, 2001: 50 m transect (Flowering ramet density/m2) Pinus banksiana 60

1986 50 2004

20 Percentage of stems

0 < 0.5 dm > 0.5-1.0 dm > 1.0-2.0 dm > 2.0 dm Size class distribution

Change in Vegetation Structure 70

60 1986 50 2004

Percent cover 20

0 Savanna P.banksiana Prairie Rhamnus

Clarke & Pine Pinus banksiana

50 m transect (total population cencus by size class, vegetation structure by % cover) 20

Hypericum kalmianum

10 % frequency 5

0 1986 X2 P < 0.001 2004

Clarke & Pine Hypericum kalmianum

South Baseline: 500 m transect (% frequency/m2)

35 Rhus aromatica var. arenaria 30

% frequency 10

0 1986 X2 P = 0.055 2004

Clarke & Pine Rhus aromatica var. arenaria

South Baseline: 500 m transect (% frequency/m2)

18 Vegetation Structure 16

(+se) 10 2

Species/1-m 4

0 Woody Graminoid Forb Total Native 1976 WM 2004 WM 1976 DM/M 2004 DM/M

Woody Graminoid Forb Total Native Habitat Richness Richness Richness Richness Wet-mesic (WM) ANOVA Probability 0.402987 0.017586 0.003147 0.0050 . Dry-mesic/mesic (DM/M) ANOVA Probability 0.692686 0.226097 0.041573 0.1844 .

South Baseline: 500 m transect (species density/m2 1986: 99 plots, 2004: 50 plots)

50 Rhamnus frangula 45

% frequency 10

1986 X2 P = 0.004 2004

Clarke & Pine Rhamnus frangula

South Baseline: 500 m transect (% frequency/m2) Appendix III. Hoosier Prairie Nature Preserve Species Location & Q.# Plot Data Change

Carex conoidea NIPSCO ROW #1 1 m-sq, 5 m x 6m genet size & density, 1/4 m-sq frequency unknown Corydalis sempervirens Sand savanna #1,2 8 linear transects (15-34 m) linear frequency present Cypripedium candidum Sand prairie #1 1 m-sq plot density, total census stable Eleocharis wolfiii Sand savanna #1 1 x 13 m transect flowering ramets/1/4m-sq stable Eriophorum angustifolium Marsh #1 5 m-sq plots presence/dm-sq decline Geranium bicknellii Sand savanna #1 1 m x 2 m genet size and density present Malaxis unifolia Sand savanna #1 200 m-sq plot total census, % flowering, density increase Platanthera flava herbiola Sedge meadow #1 1 m-sq total census, % flowering stable Polygonum caryei Sand savanna #1 14 m-sq ramet size and frequency/dm-sq unknonw Polytaenia nuttallii Sand prairie #2 100-300 m-sq total census, % flowering stable? Rhynchospora globularis recognita NIPSCO ROW #1 4 m-sq genet size and density not found Satureja arkansana Sand prairie #1 1 m-sq, 0.01 ha flowering density/1/4m-sq stable?

Cypripedium candidum 80

30 Total ramet census

0 1988 Burn 1989 Non-burn 1990 Burn 2004 Non-burn

35 Cypripedium candidum 30 Flowering Nonflowering 25

20 plot 2 15

10 Ramets/m

0 1983 1985 1986 1988 1989 1990 2004

Burn Burn Nonburn Burn Nonburn Burn Nonburn

Hoosier Prairie Cypripedium candidum Q. #1

Total census & plants/m2 plot

60 Eleocharis wolfii 50

40 2

20 % frequency per 1/4m 10

0 1986 1989 2004 x 2 P = 0.382

Hoosier Prairie Eleocharis woofii

1 m x 13 m linear transect (% frequency per ¼ m2)

100 Malaxis uniflolia 90

20 Total census & % flowering census & % Total

0 1987 1988 1989 1990 1991 2004 Total % flowering

5 Malaxis unifolia 4.5

3.5

2 3

2.5

2 Density/m 1.5

0.5

0 1987 1988 1989 1990 1991 2004 ANOVA P < 0.001

Hoosier Prairie Malaxis unifolia

200 m-sq plot (total census, % flowering, density/m2) 25 90 Platanthera flava var. herbeola 80

20 70

60 15 50

40 10

30 ramets flowering % Total census (ramets)

20 5

0 0 1986 1990 2004

Total census % flowering

Hoosier Prairie Platanthera flava var. herbeola

1 m2 plot (total census & % flowering)

300 35 Polytaenia nuttallii 250 30

25 200 20 150 15

100 % flowering 10 Total census

50 5

0 0 1986 2004 Total census % flowering

Hoosier Prairie Polytanenia nuttallii Upper: Spatial pattern in 2005 (x = conduit, * = flowering, o =non-flowering) Lower: Census and % flowering in 1986 and 2005

Satureja arkansana 250

200 2

150

100

Flowering ramets per 1/4m 50

0 1976 2004

Hoosier Prairie Satureja arkansana 1986: 1 m2 plot (flowering density per ¼ m-sq in three patches) 2004: 0.01 ha plot (flowering density per ¼ m-sq in three patches)

Appendix IV. Indiana Dunes Nature Preserve Species Location & Q.# Plot Data Change

Botrychium matricariaefolium Sand Forest (Trail 2) #1 2 m x 550 m total census decline Botrychium oneidense Sand Forest (Trail 2) #1 2 m x 550 m total census stable Cakile edentual Lakeshore #1,2,3,4,5 2 m x 50 m density/10m-sq decline? Carex folliculata Flatwoods #1 2 m x 40 m % frequency/m-sq stable Carex pedunculata Sand Forest #1 2 m x 16 m ramets/m-sq stable Carex seorsa Flatwoods #1 2 m x 40 m % frequency/m-sq stable Chimaphilla umbellata Sand forest 2 m x 24 m ramets/m-sq decline Cirsium pitcheri Blowout #1 1/10 ha Population structure decline Cypripedium parviflorum Flatwoods #1 census ramets/genet decline Epigea repens Trail 9 #1 1 m-sq ramet density decline Epigea repens Mt. Jackson #2 1 m-sq ramet density decline Epigea repens Dune Creek Bluff #4 6 m x10 m frequency/1/4 m-sq stable? Euphorbia polygonifolia Lakeshore #1,2,3,4,5 2 m x 250 m genets/m-sq decline Juniperus communis depressa Foredune #1 1/10 ha size, pop. structure stable Lathyrus japonicus Foredune #1 10 m-sq ramet density decline Melampyrum lineare Dune Creek #2 60 m-sq census, density/m-sq decline Melampyrum lineare Flatwoods #3 15 m-sq census, density/m-sq decline Millium effusum Sand Forest Trail #2 2 m x 2000 m census, density/50 m-sq stable Oryzopsis asperifolia Sand Forest #1 160 m-sq demographic decline? Platanthera clavellata Flatwoods #1 2 m x 40 m % frequency/m-sq stable Platanthera flava herbiola Flatwoods #1 1 m-sq genet size & density unknown Poa alsodes Sand Forest Trail #2 2 m x 2000 m census, density/50 m-sq increase? Polygala paucifolia (white) Mt. Jackson Q#1,2 1/20 ha, 1 m x 24 m density/1/4 m-sq, % flwrng decrease? Polygala paucifolia (pink) B. house trail Q#3,4 0.016 ha, 5m-sq density/1/4 m-sq, % flwrng stable Potentilla anserina Blowout #1 1 m-sq ramet size & density decline Pyrola elliptica Sand Forest 12 m-sq density/m2 stable Pyrola rotundifolia Dune Creek Bluff 240 m-sq % frequency/1/4 m-sq decline Salix syrticola Foredune 2 m x 30 m transect density/m2 decline Stipa avenacea Sand Forest poulation census stable

30 Botrychium matricariaefolium 25

10 Number of plants

0 1984 1986 1990 2004

Indiana Dunes Botrychium matricariaefolium

20 m x 550 m linear transect (total census)

30 Botrychium oneidense 25

10 Number of plants

0 1984 2004

Indiana Dunes Botrychium oneidense

20 m x 550 m linear transect (total census) 50 Cakile edentula 1985 40 1988 1989 30 2004 2

20 Plants/10-m

0 TR1 TR2 TR3 TR4 TR5

Indiana Dunes Cakile edentula

250 m x 2 m linear transect (density/10-m2 plot in five 50 m units)

Carex folliculalta 35

15 Plot frequency 10

0 1986 2004 X2 P = 0.345

Indiana Dunes Carex folliculata

2 m x 40 m linear transect (frequency/m2 plot in 20plots)

7 Carex pedunculata

4 2

3 Ramets/m

0 1989 2004 t-test P = 0.483

Indiana Dunes Carex pedunculata

2-m x 16-m transects (ramet density/m2)

Carex seorsa 90 80 70 60 50 40 30 Plot frequency 20 10 0 1986 X 2 P = 0.178 2004

Indiana Dunes Carex seorsa

2 m x 40 m linear transect (frequency/m2 plot in 20 plots) Indiana Dunes 100

90 Cirsium pitcheri 80

1/10 ha plot cencus 20

0 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 Year

Indiana Dunes Cirsium pitcheri Upper: Big Blowout 1/10 ha plot census data. Lower: Big Blowout GPS locations Juniperus communis 9

3 Mean area in sq meters 2

0 1986 t-test P = 0.210 2004

Juniperus communis 90

80 1986 70 2004 60

% of plants 30

0 Female Male Juvenile X 2 P = 0.004

Indiana Dunes Juniperus communis

1/10-ha plot (mean plant size, population structure) Melampyrum lineare 200 180 160 140 120 100 80 Total census 60 40 20 0 1986 2004

Melampyrum lineare

2 8

Density per m 4

0 1986 2004 t-test P =0.328

Indiana Dunes Melampyrum lineare (Quadrat # 2) , Dune Creek Bluff 60-m-sq plot (total census and ramet density/m2) 250 Melampyrum lineare 200

150

100 Total census

0 1986 1989 2004

Melampryum lineare 20

Plants/m2 10

0 1986 1989 2004 ANOVA P < 0.001

Indiana Dunes Melampyrum lineare (Quadrat # 3), Flatwoods 15-m-sq plot (total census and ramet density/m2)

225 Milleum effusum 200

175

150

125

100

Total census 75

0 1986 2004

6 Milleum effusum 5

4 2

3 Plants/50m 2

0 1986 t-test P = 0.393 2004

Indiana Dunes Milleum effusum Trail #2 Sand Forest 2 m x 2000 m transects (total census & plants/50-m-sq.)

40 Platanthera clavellata 35

y 25

15 Plot frequenc

0 1986 2004 X 2 P = 0.730

Indiana Dunes Platanthera clavellata, Flatwoods

2 m x 40 m linear transect (frequency/m2 plot in 20plots)

350 Poa alsodes 300

250

200

150 Total census

100

0 1986 2004

14 Poa alsodes 12

2 8

6 Plants/50m

0 1986 2004

Indiana Dunes Poa alsodes Trail # 2 Sand Forest 2 m x 2000 m transects (total census & plants/50-m-sq.)

150 135 Polygala paucifolia 120 1976 105 2004 2 90 75 60 45 30

Total density/1/4m 15 0 Ramet density (pink) Ramet density (white)

t-test P = 0.236 Population t-test P =0.006

50 Polygala paucifolia (white)

40 1976

30 2004

20 & % flowering 2

Density/1/4m 0 Flowering density % flowering

t-test P = 0.828 t-test P < 0.001

50 Polygala paucifolia (pink)

40 1976 30 2004 20 & % flowering & %

2 10

0 Flow ering density % flow ering Density/1/4m t-test P = 0.017 t-test P < 0.001

Indiana Dunes Polygala paucifolia White: Pink: Density per ¼ m-sq

16 Pyrola elliptica 14

2 10

Stems/m 6

0 1986 2004 t-test P = 0.516

Indiana Dunes Pyrola elliptica 12 m2 plot (stems/m2)

35 Pyrola rotundifoloia 30

25 2

15 Frequency/1/4 m 10

0 1986 X 2 P < 0.001 2004

Indiana Dunes (Dunes Creek Quad #_) Pyrola rotundifolia 240 m2 plot (stems/1/4m2)

14 Salix syrticola 12

2 8

6 Stems/m

0 1986 1990 2004

Indiana Dunes Salix syrticola 2 m x 30 m transect (stems/m-sq)

10 flowering sterile Stipa avenacea 8

4 Total census

0 1986 2004 X 2 P = 0.213

Indiana Dunes Stipa avenacea

Total population census Appendix V. Tolleston Ridges-Gibson Woods Nature Preserve. Species Location & Q.# Plot Data Change

Carex aurea Mesic prairie 6 m-sq plots genet size & density decline Carex richardsonii Dry-mesic prairie 2 m x 26 m transect density m-sq decline Cypripedium parviflorum Sand savanna 1/10 ha plot census, plant size decline Cypripedium reginae Sand savanna 5 demographic plots census, plant size, canopy cover variable

80 80 Dry-mesic Prairie - 1986 Dry-mesic Prairie - 2004 70 70

60 60 Alien Native 50 50 Native 40 40

30 Relative abundance 30 Relative abundance Relative 20 20

10 10

0 0 Graminoid Woody Forb Graminoid Woody Forb

Tolleston Ridges Change in vegetation structure in dry-mesic prairie habitat for Carex richardsonii.

4 10 Forbs 3.5 1986 9 8 3 2005 7 2.5 6 2 5 1.5 4 Species/sq-m Species/sq-m 3 1 2 0.5 1 0 0 Woody (P < 0.001) Graminoid (P = 0.05) 1986 P = 0.01 2005

Tolleston Ridges Above: change in native vegetation structure in mesic savanna habitat for Cypripedium reginae Species level changes in mesic sand savanna at Tolleston Ridges

Plot frequencies in mesic savanna at Tolleston Ridges for declining species that are often associated with tallgrass prairie.

Species 1986 2005 Aster azureus 17.24 Commandra richardsonii 17.24 7.14 Coreopsis tripteris 17.24 Desmodium canadense 6.90 Euphorbia corrolata 27.59 7.14 Lithospermum caroliniense 10.34 Monarda fistulosa 10.34 Pedicularis canadensis 20.69 3.57 Andropogon gerardii 10.34 Andropogon scoparius 13.79 Calamagrostis canadensis 13.79 Sorghastrum nutans 6.90 Salix humilis 13.79

Increasing plot frequencies of dominant species in mesic savanna at Tolleston Ridges.

Species 1986 2005 Equisetum arvense 44.83 64.29 Aralia nudicaulis 24.14 57.14 Amphicarpa bracteata 17.24 53.57 Smilacina stellata 48.28 53.57 Cornus racemosa 10.34 53.57 Podophyllum peltatum 6.90 46.43 Smilacina racemosa 20.69 46.43 Symplocarpus foetidus 10.34 39.29 Parthenocissus quinquefolia 20.69 39.29 Vitis riparia 37.93 39.29 Quercus ellipsoidalis 35.71

Cypripedium calceolus var. parviflorum 15 400 1400 Genets-1986 Cyprepedium calceolus Genets-2004 350 10 var. parviflorum 1200

300 1000 5 Genets/ha 250 Ramets/ha 800 0 0 5 10 15 20 25 30 200 600 Ramets/ha -5 Genets/ha 150 400 100 -10 Meters north (+) and south (-) of baseline 50 200 -15 0 0 1986 2004 -20

Tolleston Ridges. Cypripedium calceolus var. parviflorum. 1/10 ha plot (genet locations, genet and ramet density).

Cypripedium reginae census 60 140 Cypripedium reginae C anopy Cover

120 50

100 40

80 30 60 20

Annual census 40

20 10 Mean % canopy openness

0 0 1992 1993 1994 1996 1999 2004 2005 1995 1999 2004 2005

Tolleston Ridges. Cypripedium reginae. Five demographic plots (combined population census and % canopy openness).