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Hybridization Between Native And Introduced In And The Pacific Northwest Lindsay V. Clark and Marie Jasieniuk, Department of Sciences, University of California – Davis. Contact: [email protected] R. ursinus and R. fruticosus

Principal Coordinate Analyses of genetic distances 0.6 4,6a between individuals. 7 Introduction and Objectives Distances are based on nuclear microsatellite 10 5c

TCM genotypes. Chloroplast haplotypes are indicated by 0.4 8 Many of Rubus worldwide are invasive, RSB 5a KVY colors and symbols. 13 KLS 11 12 including the apomictic R. armeniacus in • Blue: “uncut” trnK allele, indicative of R. ursinus as

CMW 0.2 Western North America. Given that hybridization well as hybrids and maternally derived from PC2 frequently contributes to invasiveness in , FCR R. ursinus. • Red: individuals with the “cut” trnK allele that 0.0 ! our objectives are to CSR morphologically appear to be in the R. fruticosus SRP complex. 2 3 • Identify spontaneous hybridization events DMU • Magenta: individuals with the “cut” trnK allele that -0.2 1 between native and introduced Rubus or among KRC morphologically do not appear to be in the R. BWP -0.6 -0.4 -0.2 0.0 0.2 0.4 0.6 DNV fruticosus complex. PC1 introduced Rubus species that could potentially KNLCFX • Black: missing data at trnK locus. ARP PLV R. fruticosus SFD CTF • Each symbol indicates a different ndhF allele. lead to new invasive forms, and CSP • Determine whether invasive populations are all MGHGHR GST Key to circled clusters: WSH 1) R. ursinus 0.4 one clone of R. armeniacus or are derived from CMB 7 PAR 2) R. ursinus x armeniacus hybrids at CSP

other species in the European R. fruticosus SDW 0.2 COP 3) R. armeniacus complex. 4) R. ulmifolius in ARP 5b

PC2 8

5a) R. laciniatus in FCR and PLV 0.0 5b) R. laciniatus in FCR and PLV, R. armeniacus-like 6a Map of field collection sites in CMW and KRC 5c) R. armeniacus-like in CMW and KRC -0.2 9 6a) R. ulmifolius in MCD, R. armeniacus-like in FCR 3

Materials and Methods -0.4 6b) R. ulmifolius in MCD 4 • Leaf material for DNA extraction was collected from 592 individuals 6c) R. armeniacus-like in FCR 7) R. armeniacus-like in FCR, CSR, CMW 0.0 0.5 1.0 across California, , and . 8) ‘Navajo’ thornless PC1 • All individuals were genotyped with eight nuclear microsatellite 9) R. armeniacus-like in ARP, CFX, DVS R. ursinus and non-fruticosus non-ursinus 10) markers and two chloroplast markers (trnK and ndhF). 11) ‘Siskiyou’ blackberry 13 • Genetic distances between individuals were calculated based on 12) R. armeniacus-like in DVS, CSR 1 13) R. ursinus-like in BWP, COP 0.2

distances between alleles, using the method of Bruvo et al. (Mol Ecol 14) R. pensilvanicus in BWP 13: 2101) for mixed polyploid systems, implemented in R. 15) R. parviflorus in KRC and SRP 16a 16a) R. leucodermis, cultivated , , 0.0 • For 471 individuals with complete data, Principal Coordinate unidentified species in TCM, R. ursinus in KRC, R. PC2 14 Analysis was used to identify distinct genetic groups and hybrids. allegheniensis, R. oderatus, 10

16b) R. leucodermis, cultivated raspberries, -0.2 2 Rubus native and naturalized to the West Coast, obtained in field collections unidentified species in TCM, R. allegheniensis, R. ! Subgenus Rubus (blackberries) oderatus, Marionberry 16c) cultivated raspberries, Marionberry 15

Sect. Ursini -0.4 R. ursinus (Pacific blackberry, native) 16d) R. leucodermis, unidentified species in TCM, R. Sect. Rubus (R. fruticosus agg.) allegheniensis, R. oderatus -0.8 -0.6 -0.4 -0.2 0.0 0.2 R. armeniacus (Himalayan blackberry, Europe) PC1 R. laciniatus (Cutleaf evergreen blackberry, Europe) R. fruticosus and non-fruticosus non-ursinus R. ulmifolius (Elm-leaf blackberry, Europe) 14

Sect. Arguti 0.4 R. pensilvanicus (Pennsylvania blackberry, Eastern North America) 15,16b Subgenus Ideaobatus (raspberries) 3 R. leucodermis (blackcap , native) 0.2 9 R. spectabilis (salmonberry, native) 8 Subgenus Anaplobatus 0.0 5a R. parviflorus (thimbleberry, native) PC2

-0.2 4,6b Other Rubus included in study 6c

• R. allegheniensis – Subgen. Rubus Sect. Allegheniensis, Eastern North America A -0.4 • R. oderatus – Subgen. Anaplobatus, Eastern North America

• R. idaeus (red raspberry) – cultivated raspberry originating in Europe – cultivars ‘Cuthbert’, -0.6 5c,7 ‘Chilliwack’, ‘Cascade Delight’, ‘Autumn Britten’ • – R. ursinus x idaeus -0.4 -0.2 0.0 0.2 0.4 0.6 0.8 1.0 • Boysenberry – Loganberry x blackberry x raspberry PC1 • – Loganberry x raspberry x non-fruticosus non-ursinus • Marionberry – R. armeniacus x R. ursinus x Loganberry x Ollallieberry • Thornless Marionberry

• Tayberry – blackberry x raspberry 0.2 • ‘Navajo’ thornless blackberry 15 16c 14 • ‘Siskiyou’ blackberry – Ollallieberry x Boysenberry B

• ‘Chester’ thornless blackberry 0.0

• ‘Silven’ blackberry PC2

Conclusions -0.2 16d • Spontaneous hybridization between the invasive R. armeniacus and common

native R. ursinus occurs rarely. -0.4

• R. armeniacus in Central and Southern California appears to be of a single C -0.4 -0.2 0.0 0.2 clonal lineage, but multiple clones are present in invasive populations in PC1 Example of morphological Acknowledgements Northern California and the Pacific Northwest, potentially derived from intermediacy of hybrids Stella Hartono, Anna Sherwood, Annabelle Kleist, A) R. armeniacus. B) R. ursinus. C) hybridization with R. laciniatus, R. ulmifolius, and/or other species. Jeffrey Firestone, and Noor-ul-ain Noor for lab and R. ursinus x armeniacus. • populations that are maternally derived from R. ursinus are present in field assistance; Cosumnes River Preserve, Caswell Memorial State Park, American River Cherry Bidwell Park (BWP) and Coal Oil Point Reserve (COP). These may be hybrids Company, and Pacific Star Gardens for allowing us with R. pensilvanicus or a species not included in this study. to collect samples; Coal Oil Point Reserve, Paradise Reserve, and Sedgwick Reserve for mailing samples • R. ursinus, which is thought to be an ancient hybrid between a blackberry and to us; Kristina Schierenbeck, Susan Lambrecht , and raspberry, hybridizes freely with native and cultivated raspberry species. Ramona Robison for leads on interesting blackberries; and Jastro-Shields Research Award • Popularly cultivated hybrid Rubus has not contributed to invasive populations. and Plant Sciences Departmental Research Assistantship for funding.