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Thesis Capstone: Plant Assemblages in Constructed and Natural Vernal Pools in New York State
- 1. Vernal pools of the northeastern US: Plant assemblages and environmental conditions of constructed and natural ephemeral pools in New York State Jaime B. Jones 18 April 2014
- 2. Outline • Overview • Objectives • Methods – Sampling design – Data collection • Results & Recommendations – Abiotic factors – Vascular plant cover and richness – Species composition – Propagule bank – Influences on species composition • Framework for conceptual model • Future research
- 3. Vernal pools in the northeastern US • Small, isolated, temporary wetlands (Colburn 2004) • Amphibian breeding habitat • Gamma diversity (Colburn 2004) • Plant species of concern (Comer et al. 2005) – Carex lupuliformis (S2), Carex typhina (S1), Carex buxbaumii (S2) – Scirpus ancistrochaetus: “northeastern bulrush” (E) • Vernal pool losses conserve, restore, construct Natural pool Created pool (circa 1970)
- 4. Upper Susquehanna Coalition • Network of 19 soil & water conservation districts • Built 300+ vernal pools between 2003-2010 u-s-c.org
- 5. Upper Susquehanna Coalition Schuyler, Chemung, & Tompkins Co., NY u-s-c.org
- 6. 2007 2008 2006 2003 2006 Study sites • Eight properties • Randomly selected pools, n = 77 • 60 “new” constructed pools (2003 – 2008) (17 open, 43 forest) • 10 “old” constructed pools (~1970) • 7 Natural pools
- 7. 1. Characterize & compare constructed & natural vernal pools (CVP & NVP) Objectives 2. Describe influence of selected environmental factors on vascular plant composition, richness, & cover 3. Recommend pool designs and management strategies to facilitate establishment of desired plant species
- 8. Methods
- 9. Environmental data Abiotic • Pool age • Light (PAR) – Percent full sun (mean, range) • Depth & area – max, min, residual (%) • Specific cond, pH, temp. • Margin slope • Soils (n=30 CVPs) – Bulk density – Organic matter Biotic • Total basal area • Tree species richness • Coarse woody debris • Bryophyte cover • Terrestrial vegetation – Cover by species – Total cover – Species richness •
- 10. Vernal pool vascular plant data • Cover by species (%) • Total cover (%) • Species richness
- 11. Propagule bank study • Composite soil samples, constructed pools (n=15) • Flooded & mesic treatments • Seedling emergence method (van der Valk and Davis 1978, Smith and Kadlec 1983, Haukos and Smith 1993) Flooded treatments Mesic treatments
- 12. Results: Abiotic factors Pool type Mean pH Const., new open 6.13 ± .44 Const., new forest 5.65 ± .57 Const., old forest 5.58 ± .54 Natural, forest 5.25 ± .52
- 13. Light availability (%PAR penetration) of CVPs and NVPs Whiskers extend to 5th and 95th percentiles All differed significantly except NVPs and new forest CVPs. n=10 n=17 n=43 n=7
- 14. Residual depth (%) of CVPs and NVPs Whiskers extend to 5th and 95th percentiles Residual pool Significant differences: Old CVP & new CVP NVP & new CVP NO significant differences: NVP & old CVP New open CVP & new forest CVP
- 15. Average margin slope of CVPs and NVPs Slope Old CVPs New CVPs NVPs
- 16. Soils Bulk density (g/cm3) in CVP margin soils Whiskers extend to 5th and 95th percentiles ------------Root-restrictive Db in clayey soil (n=30 USC-created pools) Organic matter (%) in CVP margin soils Whiskers extend to 5th and 95th percentiles 3.3 to 21.95% 0.57 to 1.53 g/cm3
- 17. Results: Vascular plant assemblages
- 18. Mean Mean total total cover cover Const.(old) Const.(new)open Const.(new)forest Natural Origin
- 19. MMeeaann sp teociteas lr icchonevses r Const.(old) Const.(new)open Const.(new)forest Natural Total of 158 vascular plant species
- 20. Results: Influences on cover and species richness
- 21. Modeling total cover and species richness (constructed pools) Generalized Linear Mixed Models – Main effects: selected environmental factors – Random effect: property – Selecting top candidate models: Akaike Information Criteria (AIC)
- 22. Total cover Model Rank K AICc ΔAICc Wi Light pH Cond Age MaxDep MinArea Slope Total Cover 1 5 549.4 0.0 0.429 X X X X 2 4 551.1 1.7 0.180 X X X 3 6 551.7 2.3 0.134 X x X X X 4 7 552.4 3.0 0.095 X X X x X x 5 7 552.8 3.4 0.078 X x X X X x 6 7 554.0 4.6 0.043 X x X X x X Top candidate models for cover: – Light availability (+) – Pool age (+) – Specific conductance (+) – Minimum area (-) Parameters • More drawdown, more cover. Less drawdown, less cover.
- 23. Species richness Top candidate models for richness: – Light availability (+) – pH (+) – Maximum depth (+) – Specific conductance (-) Parameters Model Rank K AICc ΔAICc Wi Light pH Cond Age MaxDep MinArea Slope Species Richness 1 5 149.6 0.0 0.486 X X X X 2 6 151.6 2.0 0.183 X X X x X 3 7 153.9 4.2 0.058 X X X x X x 4 4 154.1 4.5 0.052 X X X 5 7 154.2 4.6 0.049 X X X x X x 6 4 154.2 4.6 0.048 X X X
- 24. Results: Species composition Analyses: •Multi-Response Permutation Procedure (MRPP) •Indicator Species Analysis •Life history groups
- 25. Multi-Response Permutation Procedure • Tests for difference in species composition between 2+ groups • “A” = effect size: If Then All items are identical within groups A = 1 Heterogeneity within groups equals expectation by chance A = 0 Heterogeneity within groups is greater than expectation by chance A < 0 In ecology, A > 0.3 is fairly high
- 26. Multi-Response Permutation Procedure 1. Natural- forest 2. Constructed- old, forest 3. Constructed- new, forest 4. Constructed- new, open-canopy A = 0.353 (p=0.0000000)
- 27. Indicator Species Analysis • Identifies species indicative of a priori groups • “Perfect” indicator species: – Indicator Value = 100 – Faithful – Exclusive
- 28. Indicator Species Analysis Pool Species WIS ShadeTol Native Lifespan Lifeform IV p Natural Osmunda regalis OBL Y Y PER Fern 57.1 0.0002 Carex intumescens FACW+ Y Y PER Graminoid 56.0 0.0002 Quercus rubra FACU- Y Y PER Tree regen 47.8 0.0004 Maianthemum canadense FAC- Y Y PER Forb 33.5 0.0074 Constructed, old forest Agrostis canina FACU Y Y PER Graminoid 50.0 0.0002 Galium asprellum OBL Y Y PER Forb 50.0 0.0002 Lycopus uniflorus OBL Y Y PER Forb 41.5 0.0096 Populus tremuloides -- N Y PER Tree regen 39.5 0.0300 Bidens connata FACW+ Y Y ANN Forb 35.2 0.0140 Amelanchier arborea FAC- Y Y PER Tree regen 34.6 0.0044 Leersia oryzoides OBL Y Y PER Graminoid 31.2 0.0248 Cornus amomum FACW Y Y PER Shrub 30.0 0.0024 Constructed, new open-canopy* Typha latifolia OBL N Y PER Forb 70.4 0.0002 Juncus tenuis FAC- Y Y PER Graminoid 69.1 0.0002 Juncus effusus FACW+ N Y PER Graminoid 68.1 0.0002 Potentilla simplex FACU- Y Y PER Forb 68.1 0.0002 Panicum virgatum FAC N Y PER Graminoid 64.7 0.0002 Carex scoparia FACW N Y PER Graminoid 64.0 0.0002 Euthamia graminifolia FAC N Y PER Forb 62.8 0.0004 Eleocharis ovata OBL N Y PER Graminoid 62.0 0.0004 *19 additional indicator species were identified for open-canopy pools
- 29. Plant lifeforms present in vernal pools Percentage of most abundant species (n=20) represented by various lifeforms Pool Type Percentage Graminoid Forb Fern Shrub Tree regen Constructed – Open-canopy, new 50 40 0 10 0 Constructed – Forest, new 65 25 10 0 0 Constructed – Forest, old 35 20 5 25 15 Natural- Forest 30 25 13 13 19
- 30. Results: Propagule bank study
- 31. Propagule bank germinants • Seedling density: – 63.7 - 31,433 m-2 – Positively correlated with age & light (Fenner & Thompson 2005) G. neglecta and G. uliginosum: common in seed bank, uncommon in field
- 32. Results: Environmental factors and species composition • Nonmetric Multidimensional Scaling (NMS) ordination – Pools in species space – Joint plot with environmental factor overlay – Similar pools are nearer to each other
- 33. NMS Ordination: All pools 3-D solution, stress= 19.94, instability <0.00001, explained 45.7% of variation Light and pH: Positively correlated with Scirpus cyperinus, Euthamia graminifolia, Carex scoparia, Scirpus atrovirens, Juncus effusus, Juncus tenuis, Solidago canadensis, Potentilla simplex, Typha latifolia, Eleocharis ovata, and Panicum virgatum Pool age: Negatively correlated with Onoclea sensibilis & Leersia oryzoides Δ CVP, Old forest + CVP, New forest * CVP, New open-canopy ☐ Natural
- 34. NMS Ordination: Created pools with soil data 3-D solution, stress= 16.99, instability <0.00001, explained 55.4% of variation. More compact soils: Juncus tenuis, Lolium perenne Higher organic matter: Onoclea sensibilis, Athyrium filix-femina, Arisaema triphyllum
- 35. Environmental sieves and wetland assembly Model of wetland succession (van der Valk 1981) Model of wetland assembly (Weiher and Keddy 1995)
- 36. Framework for Vernal Pool Environmental Sieve Model
- 37. Proposed species to test Environmental Sieve Model Species WIS ShadeTol Native Lifespan Lifeform Desired Carex scoparia FACW N Y PER Graminoid Y Scirpus cyperinus FACW N Y PER Graminoid Y Juncus effusus FACW+ N Y PER Graminoid Y Sparganium americanum OBL N Y PER Forb Y Typha spp. OBL N Y PER Forb N Carex lurida OBL N Y PER Graminoid Y Carex vulpinoidea OBL N Y PER Graminoid Y Eleocharis ovata OBL N Y PER Graminoid Y Scirpus ancistrochaetus OBL N Y PER Graminoid Y* Scirpus atrovirens OBL N Y PER Graminoid Y Lonicera morrowii - Y N PER Shrub N Carex leptonervia FACW Y Y PER Graminoid Y Cornus amomum FACW Y Y PER Shrub Y Carex intumescens FACW+ Y Y PER Graminoid Y Ilex verticillata FACW+ Y Y PER Shrub Y Bidens cernua OBL Y Y ANN Forb Y Osmunda regalis OBL Y Y PER Fern Y Lycopus uniflorus OBL Y Y PER Forb Y Glyceria striata OBL Y Y PER Graminoid Y Leersia oryzoides OBL Y Y PER Graminoid N
- 38. Future Research • Env. sieves- soil texture, leaf litter • Spatial analysis - Estimated dispersal distances (Tamme et al. 2014) • Successional processes • Assembly rules- seed arrival history, competition, environment • Scirpus ancistrochaetus
- 39. Northeastern bulrush conservation • Scirpus ancistrochaetus • Federally listed: Endangered • One population in NY: Steuben Co Arthur Haines
- 40. Northeastern bulrush: Ecology • Small, open wetlands • Shallow, fluctuating water • Vernal pools, wetland & pond margins • Sandy substrates (Center for Plant Conservation) Tess Feltes Steve Young
- 41. Northeastern bulrush: Future research • Survey for presence of S. ancistrochaetus in potentially suitable habitat near Steuben Co. Steve Young • Effect of environmental factors & interspecific competition • Inbreeding depression in isolated populations • Seedbank persistence- soil transplants? New impetus for VP conservation & construction
- 42. Conclusions CVPs are heterogenous, and support a variety of native FACW-OBL plant species Open-canopy CVPs differ from forested CVPs and NVPs Light availability, drawdown, depth, age, and pH: correlated with plant assemblage characteristics Environmental sieve model— frame understanding of how these environmental conditions may affect species composition Consider how design & management decisions will impact not only amphibians, but also plant assemblages
- 43. Acknowledgements Don Leopold- Major professor James Gibbs and John Stella- Committee Melissa Fierke, Steve Stehman, and Greg McGee Upper Susquehanna Coalition Edna Bailey Sussman Foundation Wetland Foundation Jessica Logan Ecolunch & 401 Illick colleagues, especially Tony Eallonardo, Kay Hajek, Jess Riddle, Patrick Raney, and Joe Gawronski-Salerno
- 45. Pool Species WIS ShadeTol Native Lifespan Lifeform Cover Const. Leersia oryzoides OBL Y Y PER Graminoid 43.3 Old Bidens connata FACW+ Y Y ANN Forb 26.6 Forest Ilex verticillata FACW+ Y Y PER Shrub 15.2 Cornus amomum FACW Y Y PER Shrub 13.6 Lycopus uniflorus OBL Y Y PER Forb 12.5 Onoclea sensibilis FACW Y Y PER Fern 11.7 Lonicera morrowii -- Y N PER Shrub 9.8 Phalaris arundinacia FACW Y PER Graminoid 9.6 Solanum dulcamara FAC- Y N PER Forb 7.7 Sparganium americanum OBL N Y PER Forb 7.0 Agrostis canina FACU Y Y PER Graminoid 5.6 Carex lurida OBL N Y PER Graminoid 4.9 Spiraea alba FAC+ N Y PER Shrub 4.7 Scirpus cyperinus FACW N Y PER Graminoid 4.0 Salix sp. -- N -- PER Tree regen 3.6 Vaccinium corymbosum FACU- N Y PER Shrub 3.5 Juncus effusus FACW+ N Y PER Graminoid 3.1 Glyceria striata OBL Y Y PER Graminoid 2.9 Acer rubrum FAC Y Y PER Tree regen 2.3 Populus tremuloides -- N Y PER Tree regen 2.3 Const. Leersia oryzoides OBL Y Y PER Graminoid 90.7 New Glyceria striata OBL Y Y PER Graminoid 88.4 Forest Sparganium americanum OBL N Y PER Forb 59.9 Juncus tenuis FAC- Y Y PER Graminoid 35.2 Scirpus atrovirens OBL N Y PER Graminoid 29.6 Impatiens capensis FACW Y Y ANN Forb 29.2 Onoclea sensibilis FACW Y Y PER Fern 24.4 Scirpus cyperinus FACW N Y PER Graminoid 22.1 Thelypteris FAC Y Y PER Fern 20.7 noveboracensis Carex leptonervia FACW Y Y PER Graminoid 19.5 Solidago canadensis FACU N Y PER Forb 19.4 Carex vulpinoidea OBL N Y PER Graminoid 18.4 Carex baileyi OBL N Y PER Graminoid 15.8 Juncus effusus FACW+ N Y PER Graminoid 14.5 Carex sp. 1 -- -- -- -- Graminoid 13.0 Carex scoparia FACW N Y PER Graminoid 12.3 Panicum virgatum FAC Y Y PER Graminoid 10.7 Lolium perenne 9.9 Euthamia graminifolia FAC N Y PER Forb 9.2 Asteraceae 1 -- -- -- -- Forb 8.9
- 46. Pool Species WIS ShadeTol Native Lifespan Lifeform Cover Const. Typha latifolia OBL N Y PER Forb 127.1 New Carex scoparia FACW N Y PER Graminoid 78.0 Open- Scirpus cyperinus FACW N Y PER Graminoid 76.3 canopy Juncus effusus FACW+ N Y PER Graminoid 75.8 Sparganium americanum OBL N Y PER Forb 68.7 Scirpus atrovirens OBL N Y PER Graminoid 65.9 Eleocharis ovata OBL N Y PER Graminoid 48.3 Typha angustifolia OBL N N PER Forb 44.2 Juncus tenuis FAC- Y Y PER Graminoid 39.2 Bidens cernua OBL Y Y ANN Forb 30.3 Agrostis gigantea -- N N PER Graminoid 22.6 Rubus sp. -- N Y PER Shrub 22.1 Carex vulpinoidea OBL N Y PER Graminoid 20.2 Bidens connata FACW+ Y Y ANN Forb 17.8 Vaccinium corymbosum FACU- N Y PER Shrub 17.1 Euthamia graminifolia FAC N Y PER Forb 16.8 Carex sp. 1 -- -- -- -- Graminoid 15.6 Solidago canadensis FACU N Y PER Forb 13.5 Potentilla simplex FACU- Y Y PER Forb 13.1 Tridens flavus UPL N Y PER Graminoid 11.1 Natural Osmunda regalis OBL Y Y PER Fern 34.9 Forest Carex intumescens FACW+ Y Y PER Graminoid 21.4 Glyceria striata OBL Y Y PER Graminoid 7.5 Ilex verticillata FACW+ Y Y PER Shrub 5.2 Lycopus uniflorus OBL Y Y PER Forb 2.7 Onoclea sensibilis FACW Y Y PER Fern 2.2 Vaccinium corymbosum FACU- N Y PER Shrub 2.2 Juncus effusus FACW+ N Y PER Graminoid 2.0 Maianthemum canadense FAC- Y Y PER Forb 2.0 Asteraceae 2 -- -- -- -- Forb 1.6 Carex lurida OBL N Y PER Graminoid 1.4 Quercus rubra FACU- Y Y PER Tree regen 1.0 Lycopus americanus OBL Y Y PER Forb 0.8 Ostrya virginiana FACU- Y Y PER Tree regen 0.6 Panicum virgatum FAC Y Y PER Graminoid 0.6 Fraxinus americana FACU Y Y PER Tree regen 0.5
Editor's Notes
- Light / pH- due to clustering of open-canopy pools on similar soils? Leaf litter? Created pool pH ranged from 4.32 – 6.82, while natural pool pH ranged from 4.42 – 6.38. Mean pH levels of created and natural pools were 5.88 0.53 and 5.25 0.61, respectively. Mean pH levels of old forest, new forest, and natural forest pools were 5.58 0.54, 5.65 0.57, and 5.25 0.52, respectively, while new open-canopy pools had a slightly higher pH of 6.13 0.44. Post-hoc Mann-Whitney tests with Bonferroni correction showed significant differences in pH between new open-canopy pools and old forest (p &lt; 0.01, r = 0.34), new forest (p &lt; 0.01, r = 0.32), and natural forest pools (p &lt; 0.01, r = 0.34). Significant differences were not detected among other pool types (α = 0.05).
- Post-hoc Mann-Whitney tests with Bonferroni correction showed significant differences among all pool groups (r = 0.31 to 0.38) except new forest pools and natural pools.
- Light / residual area- due to construction methods? (e.g. larger equipment = increased compaction = better water retention) Post-hoc Mann-Whitney tests with Bonferroni correction showed significant differences in residual area between old created pools and both types of new created pools (r = 0.38, p &lt; 0.001 for new open-canopy pools, and r = 0.31, p &lt; 0.01 for new forest created pools). Natural pools and new created open-canopy pools also differed significantly (p &lt; 0.01, r = 0.32). Significant differences in residual area were not detected between natural pools and old created pools (α = 0.05).
- significant differences in slope between old created pools and both types of new created pools (p &lt; 0.001, r = 0.38 and p &lt; 0.001, r = 0.37 for open-canopy and forest pools, respectively), and between natural pools and both types of new created pools (p &lt; 0.001, r = 0.38 and p &lt; 0.001, r = 0.37 for new open-canopy and new forest created pools, respectively).
- Post-hoc Mann-Whitney tests with Bonferroni correction: open-canopy pools differed from all other pools (r = 0.37, 0.57, and 0.41 for old forest, new forest, and natural forest pools, respectively; p &lt; 0.001). Forest pool groups (old, new, and natural) did not differ significantly from one another in total cover.
- Post-hoc Mann-Whitney tests with Bonferroni correction: new open-canopy pools differed from all other pools (r = 0.36, 0.52, and 0.46 for old forest, new forest, and natural forest pools, respectively; p &lt; 0.01). Species richness did not differ among forest pool types (new, old, or natural).
- Poisson error distributions, log link functions, and maximum likelihood criteria
- ADD COEFFICIENTS!!! Need to know if these variables are positively or negatively correlated with cover.
- ADD COEFFICIENTS!!! Need to know if these variables are positively or negatively correlated with cover.
- Non-parametric analysis. “A” Describes within-group homogeneity of sampling units (vernal pools) compared to random expectation
- Based on relative abundance & relative frequency
- Royal fern, greater bladder sedge, rough bedstraw, northern bugleweed, purple stem beggar’s ticks, broadleaf cattail, common rush,
- Out of 20 most abundant species in each type of pool. Mostly grams. More woody species in older pools.
- Marsh cudweed (G. uliginosum) Clammy hedgehyssop (G. neglecta)
- Nonparametric multivariate technique Indirect gradient analysis
- r2 cutoff for vectors was 0.25, and light was loaded on axis 1. Length of vectors indicates strength of correlation. The strongest two axes, 1 and 3, accounted for 18% and 16.7% of variation, respectively. Axis 2 accounted for 11% of variation. The strongest influences on pool plant communities were light availability and associated factors (i.e. terrestrial cover, terrestrial richness, and total basal area), pool pH, and, to a lesser extent, time since pool creation (age). A joint plot of axes 1 and 3 indicated that light availability, pH, terrestrial cover, and terrestrial richness were all positively correlated with axis 1 (r2 = 0.43, 0.41, 0.32, and 0.30, respectively). Total basal area was negatively correlated with these variables and negatively correlated with axis 1 (r2 = 0.37) (Figure 7). Scirpus cyperinus, Euthamia graminifolia, Carex scoparia, Scirpus atrovirens, Juncus effusus, Juncus tenuis, Solidago canadensis, Potentilla simplex, Typha latifolia, and Eleocharis ovata, all relatively shade-intolerant perennials, were positively correlated with axis 1 (r2 = 0.25, 0.18, 0.18, 0.17, 0.17, 0.16, 0.15, 0.15, 0.15, and 0.15 respectively). A joint plot of axes 2 and 3 indicated that pool age and tree species richness were positively correlated with axis 3 (r2 = 0.23 and 0.16, respectively) (Figure 7). Onoclea sensibilis and Leersia oryzoides were negatively correlated with axis 3 (r2 = 0.17 and 0.13), and Juncus tenuis was positively correlated with axis 3 (r2 = 0.13). Light was positively correlated with axis 2 (r2 = 0.15), as was Panicum virgatum (r2 = 0.13).
- Joint plots for NMS Ordination of created vernal pools with soil data (n=30) in species space. A three-dimensional solution (final stress =16.99, final instability = 0.00001) explained 55.4% of the variation. The r2 cutoff for vectors was 0.25. Length of vectors indicates strength of correlation. Species that were positively correlated with axis 3, and thus associated with pools of more compacted soil with less organic matter, included J. tenuis (path rush) and Lolium perenne (perennial rye grass, a non-native grass somties planted following pool construction) (r2 = 0.17 and 0.13, respectively). The following species were negatively correlated with axis 3, and were thus associated with pools of lower bulk density and higher organic matter: Onoclea sensibilis (sensitive fern), Athyrium filix-femina (lady fern), and Arisaema triphyllum (jack in the pulpit)
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