Jaime B. Jones18 April 2014

Vernal pools of the northeastern US: Plant assemblages and

environmental conditions of constructed and natural ephemeral pools

in New York State

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

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)

Upper Susquehanna Coalition

• Network of 19 soil & water conservation districts

• Built 300+ vernal pools between 2003-2010

u-s-c.org

Upper Susquehanna Coalition

Schuyler, Chemung, & Tompkins Co., NY

u-s-c.org

2007

2008 2006

2006

2003

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

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

Methods

Environmental dataAbiotic• 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

•

Vernal pool vascular plant data

• Cover by species (%)• Total cover (%)• Species richness

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

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

n=43n=17n=10 n=7

Light availability (%PAR penetration) of CVPs and NVPsWhiskers extend to 5th and 95th percentiles

All differed significantlyexcept NVPs and new forest CVPs.

Residual pool

Residual depth (%) of CVPs and NVPsWhiskers extend to 5th and 95th percentiles

Significant differences: Old CVP & new CVPNVP & new CVP

NO significant differences: NVP & old CVPNew open CVP & new forest CVP

Slope

Average margin slope of CVPs and NVPs

Old CVPs New CVPs NVPs

Soils

------------Root-restrictive Db in clayey soil

(n=30 USC-created pools)

3.3 to 21.95% 0.57 to 1.53 g/cm3

Organic matter (%) in CVP margin soilsWhiskers extend to 5th and 95th percentiles

Bulk density (g/cm3) in CVP margin soilsWhiskers extend to 5th and 95th percentiles

Results: Vascular plant assemblages

Mean total cover

NaturalConst.(new)openConst.(old) Const.(new)forest

Origin

Mean total cover

Mean total coverMean species richness

NaturalConst.(new)openConst.(old) Const.(new)forest

Total of 158 vascular plant species

Results: Influences on cover and species richness

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)

Total cover

Top candidate models for cover:– Light availability (+)– Pool age (+)

– Specific conductance (+)

– Minimum area (-)• More drawdown, more cover. Less drawdown, less cover.

Parameters 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

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

Results: Species composition

Analyses:

•Multi-Response Permutation Procedure (MRPP)

•Indicator Species Analysis

•Life history groups

Multi-Response Permutation Procedure

• Tests for difference in species composition between 2+ groups

• “A” = effect size:

In ecology, A > 0.3 is fairly high

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

1. Natural- forest2. Constructed- old, forest3. Constructed- new, forest4. Constructed- new, open-canopy

A = 0.353 (p=0.0000000)

Multi-Response Permutation Procedure

Indicator Species Analysis

• Identifies species indicative of a priori groups

• “Perfect” indicator species: – Indicator Value = 100

– Faithful

– Exclusive

Indicator Species AnalysisPool Species WIS ShadeTol Native Lifespan Lifeform IV p

Nat

ural

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

Con

stru

cted

, old

fo

rest

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

Con

stru

cted

, new

op

en-c

anop

y*

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

Plant lifeforms present in vernal pools

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

Percentage of most abundant species (n=20) represented by various lifeforms

Results: Propagule bank study

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

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

NMS Ordination: All pools3-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 virgatumPool age: Negatively correlated with Onoclea sensibilis & Leersia oryzoides

Δ CVP, Old forest

+ CVP, New forest

* CVP, New open-canopy ☐ Natural

NMS Ordination: Created pools with soil data3-D solution, stress= 16.99, instability <0.00001, explained 55.4% of variation.

More compact soils: Juncus tenuis, Lolium perenneHigher organic matter: Onoclea sensibilis, Athyrium filix-femina, Arisaema triphyllum

Environmental sieves and wetland assembly

Model of wetland succession (van der Valk 1981)

Model of wetland assembly (Weiher and Keddy 1995)

Framework for Vernal Pool Environmental Sieve Model

Proposed species to test Environmental Sieve ModelSpecies 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

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

Northeastern bulrush conservation• Scirpus ancistrochaetus• Federally listed: Endangered• One population in NY:

Steuben Co

Arthur Haines

Northeastern bulrush: Ecology

• Small, open wetlands • Shallow, fluctuating water• Vernal pools, wetland & pond margins• Sandy substrates (Center for Plant Conservation)

Tess Feltes

Steve Young

Northeastern bulrush: Future research• Survey for presence of S. ancistrochaetus in potentially

suitable habitat near Steuben Co.

• Effect of environmental factors & interspecific competition

• Inbreeding depression in isolated populations

• Seedbank persistence- soil transplants?

New impetus for VP conservation & construction

Steve Young

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

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

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

noveboracensis FAC Y Y PER Fern

20.7

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

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