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Camassia Populations (Agavoideae, Asparagaceae)

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Camassia Populations (Agavoideae, Asparagaceae) Linking floral anthesis, pollination, and species boundaries in Camassia populations (Agavoideae, Asparagaceae) Willamette University, Salem, OR Susan Kephart and Sebastian Mortimer Pollination & Species Boundaries: What we know... • Spatio-temporal variation influences plant-pollinator interactions, affecting species divergence and diversity (Burkle & Alarcón 2011) • Pollinator-mediated isolation is clearly evident when floral cues differ greatly (e.g., Mimulus, Penstemon) but less often explored in flowers accessible to generalist pollinators (Kephart & Theiss 2004; Kimball 2008) • Plant-pollinator interactions, abiotic, & historical factors can shape both floral traits and phenological timing, e.g., via selection against hybrids and/or competition for pollinator services (Stone et al. 1996). Rationale: Why Camassia? 1) Floral Traits & Speciation • Variable floral display & duration • Dynamic species boundaries w. hybrid zones (Uyeda & Kephart 2006) 2) Ecology-Conservation Roles • Diverse pollinators; Key nectar resource (Parachnowitsch & Elle 2005; Sultany et. al 2007) • Tractable data for climate change, B. Moissett pollinator decline • Habitat loss: oak savanna & wetland meadows (Wilson 1998) J. Reveal Visitors: Diverse Species & Functional Groups Hymenopteran & dipteran dominated system Social bees Flies S. Pollock A. Kotaich H. Wisch J. Ascher B.Ezray Bombus sp Xylocopa sp Apis mellifera Syrphidae Bombylidae Solitary bees D. Ditchburn K. Hillig M. Jacobson S.Cain H.Wisch Andrendidae Halictidae Megachilidae BUT ... Is that all? Visitors: Diverse Species & Functional Groups But..... S. Pollock Geography Disparity: 6 sp., 15 putative taxa Western region: putative origin & plexus of diversity (Gould 1942) NW Study Sites: Camassia Key Q’s: Geography & Pollination • Q1: Do pollinator faunas on camas differ by geography or species? • Q2: Do generalist pollinators vary in visitor abundance & effectiveness on sympatric Camassia? • Q3: How does floral anthesis differ in diurnal vs vespertine populations? • Q4: What are the implications for species conservation & future study? Pollinator Faunas: Geography vs Species Q1: Do pollinator faunas on camas differ by geography or species? Predict: functional group similarity more evident by species vs geography Test: Allopatric populations: Within vs between species comparisons Capture data: CAQU breviflora Sagehen Meadow Photo credit: S.Datwyler Within species: Allopatric populations, OR &CA C. quamash breviflora C. quamash breviflora Oregon California P > 0.25, df = 4, X2 = 1.5 No differences in pollinator composition in different geographic regions Briana Ezray WU Common geography: Allopatric Species, CA P < 0.05, df = 4, X2 = 10.22 Major visitors varied signficantly by camas species in same geographic region Common geography: Allopatric subspecies C. quamash breviflora C. quamash maxima E. Oregon W. Oregon P > 0.15, df = 4, X2 = 6.6 No significant differences in capture frequency Sympatric Pops: Insect Visitation, Effectiveness Q2: Do generalist pollinators vary in abundance & effectiveness on co- occurring Camassia? i.e., Is pollinator-mediated reproductive isolation unlikely? Tests: Sympatric Pops: In situ observations & manipulative expts Key parameters: Related species specific traits, i.e. varied floral displays, nectar levels, anthesis Camassia Species : Floral Displays & Traits Great camas Common camas C. quamash C. leichtlinii Focal sympatric population: Kingston Prairie, OR Phenology in Sympatric Populations CALE * CAQU * Kingston Prairie Near Meadow KP, Oregon A. Dennis, M. Sultany & multiple WU students across years: 2007-2015 Pollinators on Camassia: Methods • Pollinator Surveys (Dafni et al. 2005: Kearns & Inouye 1993) – Pollinator = alighted on flower w contact and actively foraging – Conducted over 3-4 wk flowering period – Reference collections to ID insect taxa • Belt Transects – Unispecific (2m x 60m/species) – Mixed (2m x 50m, co-flowering species) • Pollinator Observations & Expts (Dafni et al. 2005: Kearns & Inouye 1993) – 1 x 1 m plots – Varied floral density, species, time of day, season – Cut inflorescences to manipulate density – Caging experiment: checked deposition on pollen-free stigmas Temporal Variation in Pollinator Abundance Pollinator Abundance Transects: Composition of Pollinator Community 100% 90% 80% 70% 60% Other Diptera Bombyliidae 50% Other Hymenoptera Mean % Mean Bombus sp. 40% Apis mellifera Halictidae 30% 20% 10% 0% 5/16 5/17 5/20 5/20 5/28 5/31 5/31 6/3 6/13 6/13 6/14 6/14 6/17 6/17 CAQU CAQU CALE CAQU CALE CALE CAQU CALE CALE CAQU CALE CAQU CALE CAQU Adan Kotaich WU Part I: Pollinator Behavior & Effectiveness Bombylius major contacts Apis mellifera contacts anther Bombus worker gathers nectar Bombylius major contacts Halictid bee on petal after a Apis mellifera contacts anther Bombus worker gathers nectar anthers while drinking nectar. Halictid bee on petal after a as it probes for nectar. at ovary base; side approach anthers while drinking nectar. foraging bout on Camassia. as it probes for nectar. at ovary base; side approach foraging bout on Camassia. Pollinator Behavior & Effectiveness Insect Pollinator Visitation to Camas Stalks per Plot 8 C. quamash C. leichtlinii 7 t o l P / 6 d e t i s i 5 V s k l a t 4 S s a m 3 a C n a e 2 M 1 0 Apis Bombus Solitary Bees Other Diptera Hymenoptera Insect taxa Habitat & Species: Natural Meadow Visitation 0.5 CAQU 0.45 CALE 0.4 0.35 0.3 0.25 0.2 www.ubcbotanicalgarden.org 0.15 0.1 0.05 Mean # Visits Natural Camas/10 min Natural Mean # Visits 0 Near Meadow Far Meadow Habitat 2 - Insect visitation higher on CALE than CAQU (X =117.06, P <0.05) www.markshepart.net - Visitation greatest in near meadow (X2= 132.76, P <0.05) Do Abundances Vary With Admixture? Unispecific vs Interspecific Transects Camassia Pollination: What are we missing? C. quamash C. leichtlinii C. cusickii C. howellii Camassia howellii: Unusual Pollination Interesting Vespertine Pollination in Camassia... Camassia howellii: Unusual Pollination Interesting Vespertine Pollination in Camassia... Key Q’s: Anthesis Pollination mode • Q3: How does floral anthesis differ in diurnal vs vespertine populations? What are the implications for species divergence & future study? NW Study Sites: Camassia C. quamash ssp maxima Western Camassia Species C. quamash C. leichtlinii C. cusickii C. howellii Floral Anthesis: Why be Vespertine? Diurnal? • For Plants? • For Pollinators? Which mode came first? By what mechanism did pollination shift occur? Is it only present in C. howellii, and in all populations of this species? Willamette Valley: SW Oregon: C. quamash C. howellii 40 Total New Open Flowers Total New Open Flowers 35 14 30 12 25 10 20 8 15 6 10 4 5 2 0 0 DIURNAL VESPERTINE How are these patterns similar? Different? Anthesis: When do flowers open & present pollen? 3.5 3.0 C. quamash 2.5 2.0 1.5 1.0 0.5 0.0 5.0 4.5 C. howellii Ave # Stems PP 4.0 3.5 Ave # Open Flowers 3.0 Ave # New Open 2.5 2.0 1.5 1.0 0.5 0.0 0601-0900 0901-1200 1201-1500 1500-1800 1801-2100 2101-2400 Coast Range: Mt Hebo C. quamash 100% 90% 80% 70% 60% 50% 40% 30% Out of Total Buds Out of Total 20% Percent New Open FLowers New Open FLowers Percent 10% 0% www.centralcoastjournal.com//hebo Morning Afternoon Evening Opening of New flowers DIURNAL What did we learn? • Pollinators discriminate among accessible Camassia flowers w different species-level traits, but intermixed sympatric populations have similar faunas. • Generalist pollinator abundances & visitation rates differed with Camassia species, habitat, season, and floral traits. Solitary & social bees prevalent, but Bombus deposits more pollen/stigma & visits more plants in larger C. leichtlinii • Pattern not likely to reflect selection for reproductive isolation, as – insects move between species in close sympatry – pollinator assemblages varied significantly with season and habitat – additional studies show among year & site variation in pollinators w. specie • Vespertine pollination characterizes C. howellii but appears to have arisen &/or been lost multiple times during speciation and radiation of Camassia lineages Acknowledgements Acknowledgements Particular thanks to..... Many people & Agencies! !!! Willamette Students 2006-2015 Summer Teams Jim Kephart Jenny Archibald et al. Theresa Culley et al. Kathryn Theiss Lowell Ahart & the Daley’s Ed Alverson This research was made possible by: David Hays, David Wilderman City of Salem, Oregon Parks Lawrence Janeway Earthwatch Institute Vonnie & Dan Lincoln MJ Murdock Charitable Trust Chris Wagner National Science Foundation UC Berkeley Sagehen Creek FS The Nature Conservancy Willamette University SCRP Bureau of Land Management, US Forest Service Camas Team 2015... Kudos! Sage Advice ....Future Studies “any classification scheme should be recognized as an hypothesis subject to testing and modification; and finally that there is no substitute for careful natural history.... knowledge about the natural world gained through observation, experimentation, and any other means available.” (Waser et al. 2011) From Oregon to Idaho, etc! Deciphering Species Interactions: The Future Challenges Discover: What are the species? integrating multiple criteria Morphological variation Dynamic in Space & time: What are the patterns, driving forces? Prioritize: What and why? theory and application Genetic structure Ecological differentiation Geography Disparity: 6 sp., 15 putative taxa Western region: putative origin & plexus of diversity (Gould 1942) .
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