Typha Latifolia = Native • Typha Angustifolia = Believed to Be Invasive from Eurasia… Debatable • Typha X Glauca = Native Or Invasive?

• Aggressive growth results in monocultures that are hard to eradicate

• Decrease native plant biodiversity

• Effects reverberate through the food web

• Effects on ecosystem properties? www.wnps.org/invasive_species.htm

Cattails (Typha spp.) http://www.nativetreesociety.org/invasives/index_invasive.htm Midwest “T. angustifolia is probably native to Eurasia but is now established throughout much of the US. It is abundant in the Midwest, where it hybridizes with common cattail to produce the mostly sterile “hybrid cattail” (Typha x glauca).” 1) Are Typha spp. native or invasive? 2) How do we identify Typha spp.? 3) Given that Typha spp. hybridize, are the hybrids sterile?

• Concept of species gets blurred… • Consequences for restoration PROBLEM 1: Are Typha spp. native or invasive? • Typha latifolia = native • Typha angustifolia = believed to be invasive from Eurasia… debatable • Typha x glauca = native or invasive?

+ = Typha latifolia

Typha angustifolia Pollen records in Piermont Marsh, NY

European Settlement ~ 1650

Pederson et al. 2005 Shih and Finkelstein 2008

Pollen from Typha latifolia and Typha angustifolia has been present since the Holocene (~12,000 years before present) T. angustifolia and T. latifolia pollen

Tetrads Monads

Typha x glauca = dyads / tetrads / triads Identification based on pollen (Finkelstein 2003) • Typha angustifolia = monads (22.7 +/- 2.6 m), more angular • Sparganium = monads (25.3 +/- 2.7 m), more rounded Separating T. angustifolia and T. x glauca How are invasive species defined? • Non-indigenous species or strains that replace native vegetation, causing economic, environmental, and human health harm

Broader definition: includes non-native AND native species that heavily colonize an area • Morphological traits overlap between parental species and hybrids • High variability within a species

NPS/Joy Marburger http://www.nature.nps.gov/YearinReview/YIR2005/04_B.html PROBLEM 3: Are hybrids sterile?

• It depends on the hybrid…

• First-generation (F1) hybrids thought to be sterile

• Introgression may be widespread – Back-crosses to either parent are more common than previously thought, at least for Typha x glauca – Advanced generation hybrids – Hybrid swarm Problem 2 + Problem 3 = Typha spp. are a genetic headache

• Use of molecular markers (different mutation rates) – Isozymes / VNTR / AFLP / RAPD / Microsatellites / DNA Sequencing Who’s working with molecular tools?

• Isozymes – McNaughton 1965 (Stanford University) – Lee & Fairbrothers 1969, 1973; Lee 1975 (Rutgers University) – Mashburn et al. 1978 – Sharitz et al. 1980 (University of Georgia-SREL) • VNTR – Keane et al. 1999 (University of Cincinnati, OH) • AFLP – Lamote et al. 2005 (Belgium) • RAPD – Marcinko-Kuehn et al. 1999 (McMaster University, Ontario, Canada) – Snow, Selbo, Goldberg, and Wildova (Ohio State University / U. of Michigan) – Travis, Windels, and Marburger (University of New England / INDU) – Geddes and collaborators (NEIU) • Microsatellites – Tsyusko-Omeltchenko et al. 2003; Tsyusko et al. 2005 (Ukraine / U of GA-SREL) • DNA sequences – Zhang et al. 2008 (Florida Atlantic University; Typha domingensis and Typha latifolia) Molecular tools: RAPDs RAPD primer A8

Site/ L1 G3 A Ab L2 A L1 morphology

L G A L L A L Species 2 kb 1.8 kb

1 kb

Band latifolia angustifolia

1.0 kb + – 1.8 kb – + 2.0 kb – +

Diagnostic band scores from Marcinko Kuehn et al. 1999 Back-crosses are common

From Travis et al. (unpublished)

SM L A L A G B B SM

T. latifolia L T. latifolia L T. angust. A T. latifolia L From Wildova and Snow (unpublished) 1 2 3 4 5 6 7 8 9 10

http://mipn.org/Final%20YIR%202005%20Cattail%20Sleuths%2011-26-05.doc Figure. 2. DNA analysis of 10 plants from St. Croix National Scenic Riverway area. Note plant 6 has a different banding pattern, showing a backcross to T. angustifolia. All plants have both T. latifolia and T. angustifolia genetic material. Photo provided by Steve Travis, USGS. . • Is there a difference in how different Typha species affect ecosystem properties? – Plant species richness – Nutrient pools (C, N, and P) – Nitrogen transformation (denitrification, nitrogen fixation)

• Typha species identified using a complementary approach: – Morphological traits – “Ecological information” – Molecular tools L2

L1 N

G1 R G2 NATIVE

ANGUSTIFOLIA

GLAUCA

Morphological traits

LATIFOLIA

glauca

angustifolia “Ecological information”

Angustifolia Glauca

Latifolia

Restored Native (glauca)

N R L2 L1 G2 G3 G1 A

L2 L1 R

A G2 G3 G1 Molecular tools: RAPDs RAPD primer A8

Site/ L1 G3 A Ab L2 A L1 morphology

L G A L L A L Species 2 kb 1.8 kb

1 kb

Band latifolia angustifolia

1.0 kb + – 1.8 kb – + 2.0 kb – +

Diagnostic band scores from Marcinko Kuehn et al. 1999 • Genetic analyses (RAPDs) and ecological information to identify species

• 3 different species were transplanted as rhizomes: – Typha latifolia (parental species) – Typha angustifolia (parental species) – Typha x glauca (hybrid)

• Currently analyzing 100 specimens per species using molecular tools

18 days

Growth pattern supports genetic identification

P < 0.001 a P < 0.001 a a

b Number of new sprouts Mean leaf length (cm) 90 P < 0.001 R2 = 0.812

70 Mean SOM (%) 60

Site (by geographic location)

Invasion history (older to younger) 0.04 0.15 P < 0.001 P < 0.001 2 R = 0.782 R2 = 0.962 0.03 0.10 (mg/g soil) (mg/g soil) 3 0.02 4

0.05

Mean NO 0.01 Mean NH

0.00 0.00

Site (by geographic location) Site (by geographic location)

Invasion history (older to younger) 6 2.0 P = 0.002 P = 0.001 R2 = 0.718 R2 = 0.741 5 1.5

4 1.0 3

0.5 2 Nitrogen fixation (nmol/g/d) Mean denitrification (nmol/g/d)

1 0.0

Site (by geographic location) Site (by geographic location)

Invasion history (older to younger) • Not all Typha are equal – Underscores the need for proper identification using a complementary approach

• Typha species differentially affect ecosystem properties

• Soil “legacy” from invasive species may have implications for restoration – History of invasion may be critical in determining these legacies – Restoration may not be effective if soil legacies are not addressed Ross Orr 2005 • NSF (DBI 0610405 to PG) • IL-IN SeaGrant (NOAA) (co-PIs Larkin (CBG) and Tuchman (LUC)) • Carbon Scholarship (LUC) to T. Grancharova

• J. Kelly and T. Grancharova • Bryan and Paula Pickett • Bobbi Lammers-Campbell • Joy Marburger (NPS), Dan Mason (NPS), Sean Burns (GLERC intern) • Blake Anderson, Eric Appelbaum, Priscilla Andrade, Chris Calderaro, Monika Freyman, Amy Galanter, Kathi Jo Jankowski-Giefer, Jeff Kampman, Dan Larkin, Gina Lettiere, Shane Lishawa, Owen McKenna, Mark Mitchell, Michal Olszewski, Raphael Porto, Joe Schluep, Anna Sjodin, Cindy Stowell, Lane Vail, Alison Varty