Non-Local Natives and Non-Native Locals: Considerations in the Management of Rare or Endangered Species

Deborah L. Rogers Center for Natural Lands Management and AES/ Sciences, UC Davis

A nd:

Peter W. Dunwiddie (CNLM) Jennifer DeWoody (NFGEL, USDA FS) Valerie Hipkins (NFGEL, USDA FS) Structure

1. Risks of moving species (creating or maintaining non-native)

2. Risks of protecting species integrity

3. Risks of moving populations

• Four case studies

• Western US (WA, OR, CA)

• Rare, endangered plant species 1. Risks of moving species (creating or maintaining non-natives)

Case study #1: Taylor’s checkerspot and Lanceleaf (English) plantain

Butterfly status: ESA Endangered October 2013

Taylor’s checkerspot Photo: Rod Gilbert (Euphydryas editha taylori) Taylor’s checkerspot: Extant locations rangewide

• 74 known historical sites • Many were lowland prairies • 13 extant sites • 4 sites lost in South Puget Trough in 1998-2000 • Historically occurred in large numbers (>10,000) • Estimates of abundance: >500 adults at 5 sites, >100 adults at 2 sites; <100 at 4

Source: Mary Linders, WDFW Prairie and butterfly habitat restoration

• Control invasive species • Restore ecological processes (fire) • Enhance species diversity • Especially larval host and nectar sources Checkerspot larval host plants

Plantago lanceolata Plectritis congesta (non-native)

Castilleja hispida Collinsia grandiflora & C. parviflora

Castilleja levisecta

Photos: Rod Gilbert Management Challenges: Phenological fit

Mid May Late May Mid June Late June Lanceleaf plantain

• Long-established non- native plant • Has not been subject of control • Now actively planted

• Primary checkerspot host plant in many populations • May be phenologically more suitable Taylor’s checkerspot and Lanceleaf (English) plantain

Butterfly status: ESA Endangered October 2013

Threat to natives?

Source of non-native?

Rod Gilbert Plantain status: Lanceleaf plantain Taylor’s checkerspot Long-established non- (Euphydryas editha taylori) native forb (Plantago lanceolata) Case Study #2: Brewer Spruce (Picea breweriana)

• Endemic to Klamath Mountains (SW OR and NW CA) • Lives up to 900 years Brewer spruce: Projected climate niche in 2090

• Projected to disappear from Klamath Mountains by 2090

• Suitable climate moves 1500- 1700 km or more north

• Assisted colonization creates a non-native

Source: Ledig et al. 2012. A.J. Bot. 99: 2017-2030 Assisted Migration:

Creating a non-native

Risks? 2. Risks of protecting species integrity

Case study #3: Spartina foliosa and hybridization with non-natives

Distribution of invasive Spartina across one reporting region (San Mateo) ( 3. Risks of moving populations

Case Study #4: San Diego Thornmint

ilicifolia (Gray) Gray () • Annual winter herb • Protected as a threatened species federally and an endangered species by the state of California • Subject of ongoing conservation, Photos: Sarah Godfrey CNLM restoration, and mitigation efforts • Impetus for genetic study • San Diego County: 55 extant occurrences • Baja California Norte: 13 records, uncertain status • This study: 21 collections

Leaf collections

Isozyme population study: 18 loci in 3 buffer systems

Modest levels of variation (A=3.5, 85 % polymorphic) Hierarchical test of differentiation (Fst)

Model FIS FIT FST One-level: Observed 0.198* 0.322* 0.154* 20 km Permuted -0.002 1.0E-05 -9.7E-05 95% C.I. (-0.015, 0.011) (-0.011, 0.011) (-0.004, 0.005) Three-level: Observed 0.139* < 20 km Permuted 0.178 95% C.I. (0.146, 0.208) Observed 0.167 20 – 50 km Permuted 0.177 95% C.I. (0.145, 0.219) Observed 0.259* > 50 km Permuted 0.179 95% C.I. (0.122, 0.241)

“Adaptive” Genetic Variation = Common Garden Experiment

• Container experiment Laurie Lippitt, Stacy Anderson and Bryan Endress San Diego Zoo Global within species’ natural range • 55 plants from each of 5 populations (two subpops) • Outdoors, full sun • Irrigated 1000 mL per week Traits measured

Germination percent Days to first flowering Seed produced per plant Weight of seed per plant

Total plant height Plant width Number of whorls Shoot dry biomass

20 F5,251 = 13.07, P<0.0001 Phenotypic Variation

F5,261 = 8.55, P<0.0001

F5,261 = 16.67, P<0.0001

Populations from West to East Climate Differences Among Sampled Sites

• Little variation in absolute terms • Greater variance in precipitation coastal • Greater variance in temperature inland

Populations from West to East

Daymet: 36-year average Thornton et al. (2014) ORNL. Correlation between Morphological, Genetic, Climatic Differences

Spearman’s Rank Order Mean-Ppt CV-Ppt Max-Temp(Wet) CV-Temp

ρ=0.47 ρ= -0.02 ρ= -0.45 ρ=0.47 Days To Flower P<0.0001 P>0.6 P<0.0001 P<0.0001

ρ= -0.20 ρ=0.08 ρ=0.08 ρ= -0.20 No. Inflorescence P<0.0001 P>0.05 P>0.06 P<0.0001

ρ=0.03 ρ= -0.04 ρ=0.03 ρ=0.03 Biomass P>0.4 P>0.4 P>0.4 P>0.4

ρ= -0.51 ρ=0.29 ρ=0.13 ρ= -0.59 PC1 (isozymes) P<0.05 P>0.2 P>0.6 P<0.05 Implications for Management in a Changing Climate

• Increase in temperature and variability of precipitation • Range shifts in response • Messner et al. (2011) Climatic Change Evidence of polyploidy from isozymes

• 7 loci displayed more complex banding patterns (Up to 80% of samples (PGI2))

• Frequency varied among populations Interpreting Ploidy from Isozymes

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Populations from West to East Flow cytometry confirms ploidy variation

• Isozyme banding patterns suggested a minimum of three cytotypes

• DNA measurement/flow cytometry 4x provided evidence of at least two cytotypes and additional complexity

• Variation in ploidy within and among 2x populations

Diploid Tetraploid Hexaploid Summary: “Do no harm”

1. Risks of moving populations:

• Movement may be high risk • (to donor and recipient populations and/or communities) • Climate not only consideration (photoperiodism, ecology) • Ploidy – ecotypes, reproductive barriers • Loss of ability to study relationships

• Address bottlenecks with regeneration

• Conserve genetic diversity • Practice patience • Consider the soil (or canopy) seed bank • For collections, collect over time (seasons, years) • Ensure awareness of donor site history • Reduce likelihood of loss of diversity at other stages of restoration • Do you own collections or work with ‘concierge’ plant materials providers Summary: “Do no harm”

2. Risks of protecting species integrity:

• Natural changes in species range will create new ‘opportunities’ for hybridization (provides some context for considering other hybridization events) • Aggressive removal of all putative ‘artificial’ or ‘undesirable’ hybrids may erode genetic diversity and limit adaptive (and evolutionary) potential

3. Risks of moving species (creating or maintaining non-native)

• Beware of over-simplifications: use structures based on functionality, consider the nuances of each situation • Develop carefully considered management objectives: consider community level even if single-species focus; include sustainability Non-Local Natives and Non-Native Locals: Potential Risks and Benefits in the Management of Rare Species

Recognition and Appreciation

Taylor’s checkerspot case study provided by; Peter W. Dunwiddie, CNLM and University of Washington

San Diego thornmint (SDTM) study supported by:

Jennifer DeWoody (NFGEL, USDA FS) Valerie Hipkins (NFGEL, USDA FS) Bryan Endress (Oregon State University) Sarah Godfrey (CNLM) Markus Spiegelberg (CNLM) Patrick McConnell (CNLM)

SDTM Funding: CNLM and San Diego Association of Governments