Claus Holzapfel

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Clonal diversity and resistance to invasion in
remnant salt marsh patches dominated by Spartina
patens
Funded by
Claus Holzapfel Edward Kirby
with TingMin Wu, John Francois, Ildiko Pechmann
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What is Fusion Ecology?
Interactions of old and new community members

• Urban Wildlands
• Invaded wetlands
• Invaded natural Wilderness, e.g. deserts

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• Remnant Patches
• Genetic Diversity
• Common Garden Experiment
• Dynamics of the Boundary
• Physiology of Interactions
• Landscape history
• Synthesis and Future Work

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• Remnant Patches
• Whats left of the Meadowlands?
• Genetic Diversity
• Common Garden Experiment
• Dynamics of the Boundary
• Physiology of Interactions
• Landscape history
• Synthesis and Future Work

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Large and small patches of Spartina patens
highmarsh
Photo F. Artigas
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Large patch
Large gt 3,000m2 0.74 acre Small lt 200m2
0.05 acre
Small patch
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Sampled locations, large and small patches are
matched
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• Remnant Patches
• Genetic Diversity
• Are the clones in large patches genetically
  different from the small-patch clones?
• Common Garden Experiment
• Dynamics of the Boundary
• Physiology of Interactions
• Landscape history
• Synthesis and Future Work

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Spartina patens holding out against the invader
One or many clones? Are successful clones
different?
Goal genetic description of clonal structure of
patches
Ting-Min Wu, Ph.D candiate
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ISSRs (Inter Simple Sequence Repeats)

• DNA polymerase chain reaction (PCR) technique
  uses microsatellite primers designed for di- or
  tri-nucleotide repeats. The primer anchors at two
  to three random nucleotides at the 3 end
• (Gupta et al. 1994).
• Ex AGAGAGAGAGAGAGAGYC
• AGAGAGAGAGAGAGAGYA

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Electrophoresis of PCR products amplified with
UBC primers for S.patens
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Percentage of polymorphic band (PPB) 64.86

• 12 of 40 primers showed results
• Account each band for analysis

Polymorphic band
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Prepared with Popgene (version 1.32), Yeh et al.
1997), the dendrogram was prepared using Treecon
for Windows (version 1.3b), Van de Peer and De
Wachter 1994
Dendrogram generated using UPGMA (unweighted
paired group method of cluster analysis using
arithmetic average) shows genetic similarity
between clones from different sampling sites but
not within sites (i.e., between small patches (S)
and large patches (L).
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Are clones of close locations also genetically
close?
No significant correlation between genetic
distance between clones of S. patens and their
geographical distance (P 0.175, r 0.369)
genetic distance was analyzed by Canoco for
Windows 4.5 (Ter Braak, 2002).
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• Remnant Patches
• Genetic Diversity
• Common Garden Experiment
• Do clones of different locations differ in
  growth vigor?
• Are large-patch clones stronger?
• Dynamics of the Boundary
• Physiology of Interactions
• Landscape history
• Synthesis and Future Work

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Bring them (the clones) together and lets see!
Photo F. Artigas
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METHODS We planted clonal fragments (ramets)
from three Meadowlands locations with paired
large and small Spartina patens remnant patches
(and one outgroup) in a common garden design.
Growth measurement were taken during the first
growing season (2008).
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Base-growth as shown in the graph above did vary
between the four locations (ANOVA p0.007) and
between the size of patch the ramets were taken
from. Small-patch ramets consistently
show more rigorous growth than large-patch
clones (p0.008).
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• Remnant Patches
• Genetic Diversity
• Common Garden Experiment
• Dynamics of the Boundary
• Are the borderlines of large patches
  different from the ones of small patches?
• Physiology of Interactions
• Landscape history
• Synthesis and Future Work

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Large and small patches of Spartina patens
highmarsh
L
S
Photo F. Artigas
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The transect technique
Species 2
Species 1
50cm
Clonal invaders Borders - Ecophysiology - Clonal
genetics
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The overlap zone is more extensive for small
patches than for large patches. An analysis of
the relative extent of overlap (O / A B)
revealed that this differences is statistically
significant (pgt0.002, ANOVA)
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• Change from 2007 to 2008

• Large Patch
• Spartina increases its dominance strongly in
  borders of large patches (signifcant)
• Phragmites loses to small extent (but
  significantly)
• Small Patch
• Spartina decreases and Phragmites increases (but
  both not significantly)
• Patch sizes have different dynamics
  (significant interaction of transect segment and
  patch size)

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• Remnant Patches
• Genetic Diversity
• Common Garden Experiment
• Dynamics of the Boundary
• Physiology of Interactions
• How do Spartina and Phragmites compete?
• What are the competitive mechanisms?
• Landscape history
• Synthesis and Future Work

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Phragmites in the sea of Spartina problems
facing an invader
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Respiration increases from the main Phragmites
stand towards the Spartina zone
Respiration of bent Phragmites culms is lower
than of erect culm in the Spartina zone
Porometer measurements
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The Princeton Group (Peter Jaffe al.) Effects
of Spartina on sulfide build-up (and metal
sequestration). What do the roots do?
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Use of root scan technology to obtain information
of root dynamics and distribution
20cm
Secaucus High school marsh, May 2009
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• Remnant Patches
• Genetic Diversity
• Common Garden Experiment
• Dynamics of the Boundary
• Physiology of Interactions
• Landscape history
• Where large patches always large and small
  ones always small?
• Synthesis and Future Work

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The long history of the high marsh at River Bend
1930
1985
1978
1969
1959
1930
1930
1995
2001
2008
1992
Analysis Ildiko Pechmann
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The River Bend size of Spartina partens high
marsh
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Aerial imagery analysis the open marsh is
smaller than the discernible patch
Actual patch
A sign that small patches are the remnant edges
of large ones??
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How did this look like in the past?
L
S
Photo F. Artigas
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• Remnant Patches
• Genetic Diversity
• Common Garden Experiment
• Dynamics of the Boundary
• Physiology of Interactions
• Landscape history
• Synthesis and Future Work

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• Remnant Patches
• Whats left of the Meadowlands?
• A mosaic of large and small patches of Spartina
  patens high marsh.

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• Genetic Diversity
• Are the clones in large patches genetically
  different from the small-patch clones?
• YES, but close clones appear not to be closely
  related!
• Role of recruitment from seeds and/or mixing of
  clonal material?
• Current work Genetic analysis of a large amount
  of fragments (this summer)

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• Common Garden Experiment
• (a) Do clones of different locations differ
  in growth vigor?
• (b) Are large-patch clones stronger?
• YES (a) and NO (b)
• Current work Using of a part of the common
  garden as a competition experiment.
• Do small or large-patch clones win over
  Phragmites?

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Dynamics of the Boundary Are the borderlines
of large patches different from the ones of
small patches? YES, large patch borders are more
stable and are more defined. Current work
continue monitoring of transects
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• Physiology of Interactions
• How do Spartina and Phragmites compete?
• What are the competitive mechanisms?
• Physical bending indicates negative effects
  for Phragmites, light exposure is positive.
• Current work Are the root dynamics of
  Spartina correlated to the water chemistry
  (sulfides)?

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Landscape history Where large patches always
large and small ones always small? ??? Current
work aerial photo analysis ongoing
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Coauthors Edward Kirby TingMin Wu Ildiko
Pechmann John Francois TingMin Wu
Thanks to all the collaborators and
helpers MERI/Meadowlands Commisson Francisco
Artigas Ross Feltes Edward Konsevick Hackensack
Riverkeepers Rutgers Newark Huimin Man Bonder
Lab Jessica Schnell Mark June-Wells Mina
Andrews Hadas Parag Greg Burdulis Viral
Patel Sahil Wadhwa
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Restoration efforts are attempts at creating and
assembling local communities that have vanished.
Recent studies have shown the importance of using
local ecotypes of species as building blocks in
these assemblies and the need of including
information on genotypic differentiation has been
stressed. Large portions of brackish east coast
marshlands have been invaded by non-native,
European genotypes of the common reed, Phragmites
australis. As a result, only a small fraction of
the NJ Hackensack Meadowlands is now dominated by
native marsh species and only isolated patches of
Spartina patens remain. As these patches vary in
size and seem to resist encroachment by
Phragmites differentially, we are investigating
(a) whether larger patches are able to resist
invasion more than smaller patches and (b)
whether large-patch clones are better suited for
restoration efforts. In a combined approach that
includes surveys of permanent transects, common
garden transplant experiments and genetic
analysis, we are monitoring border dynamics and
assess genetic identity and performance of clones
of different patch sizes. Current results of the
ongoing project indicate that (a) border zones
between the invader and Spartina tend to be more
defined in large Spartina remnant patches than in
small patches, and (b) Spartina increases in
dominance at large-patch borders but decrease in
small-patch borders. There appears to be large
genetic differences between adjacent large- and
small-patch clones that are often more pronounced
than differences between clones of different
regions. In contrast, small-patch clones grow
faster than large-patch clones in a common garden
setting.
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• Are there genetic differences among clones of
  Spartina patens used in high marsh restoration?
• (b) Are specific clones of Spartina patens
  correlated with enhanced restoration ability
  (i.e. growth and competitive ability during
  restoration)?
• (c) Do specific clones of Spartina patens, i.e.
  those from current larger high marsh patches,
  compete more successfully with invasive
  Phragmites than clones from smaller patches
  (patch size effects)?
• (d) Can we identify specific molecular markers
  that will aid in selection of genotypes of
  Spartina patens that will be successful in
  restoration of the high marsh communities?

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Novel defenses in a sea of clones research
into the role of root exudates and the role of
clonal competition among native and exotic
grasses
John R. Quinn (1997) Fields of Sun and Grass
Claus Holzapfel, Rutgers Newark with David
Kafkewitz, Edward Kirby and Francisco Artigas
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What is Fusion Ecology?

• Formation of novel communities fused by
  coexistence of non-native and native species
• Search for coevolution among new neighbors
  resulting in easing of conflict
• Prediction of future communities and their
  ecosystem effects
• Part of Novel Ecology
• Gradient
• Pristine communities
• Novel communities
• Constructed communities

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