Applied Vegetation Science ]] (2011) 1 21

]] (2011) 1 21 SPECIAL FEATURE: VEGETATION SURVEY A phytosociological and phytogeographical survey of the coastal vegetation of western North America: beach and dune vegetation from Baja California to Alaska Manuel Peinado, Francisco Manuel Ocaña-Peinado, Juan Luis Aguirre, José Delgadillo, Miguel Ángel Macías & Gustavo Díaz-Santiago Keywords Bioclimatology; Floristic analysis; Pacific coast of North America; Plant associations; Strand; Syntaxonomy; Vegetation classes and orders Abbreviations ALC = Average Linkage Clustering; BIO = Bioclimate; MB = Macrobioclimate; PNV = Potential Natural Vegetation; RG = Relevé Group Nomenclature USDA (2010), except for the Bajacalifornian taxa (Wiggins 1980), Agave (Gentry 1978), Helianthus (Heiser et al. 1966), and Isocoma (Nesom 1991) Received 19 November 2010 Accepted 18 March 2011 Co-ordinating Editor: Milan Chytrý Peinado, M. (corresponding author, manuel.lorca@uah.es); Aguirre, J.L. (juanl.aguirre@uah.es) & Díaz-Santiago, G. (gustavo.diaz@uah.es): Cátedra de Medio Ambiente, Universidad de Alcalá, 28871 Alcalá de Henares, Madrid, Spain Ocaña-Peinado, F.M. (fmocan@ugr.es): Departamento de Estadística e Investigación Operativa, Universidad de Granada, E 18071 Granada, Spain Delgadillo, J. (jdelga@uabc.edu.mx): Facultad de Ciencias, Universidad Autónoma de Baja California, Campus de Ensenada, BC, Mexico Macías, M.Á. (mmacias@cucba.udg.mx): Departamento de Ciencias Ambientales, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, carretera a Nogales Km. 15.5, Las Agujas, Nextipac, Zapopan, JAL, Mexico Abstract Questions: What is the floristic composition of the plant communities that inhabit the beaches and dunes of the Pacific coast of North America? What are their ecological relationships in the zonal and successional gradients typical of coastal dune systems? Does climate affect the latitudinal distribution of the azonal vegetation? What other environmental factors influence their distribution on the regional or local scale? Location: Pacific coast of North America, from Baja California (23102 0 N) to Cook Inlet, Alaska (61130 0 N). Methods: A total of 1730 phytosociological relevés were obtained by sampling 279 coastal localities. In all localities, zonation was interpreted by considering transects from the shoreline inland. Through traditional phytosociological tabular classification and average linkage clustering, relevés were syntaxonomically classified. Syntaxa, supported by fidelity calculations, are described and interpreted according to their phytogeographical distribution, their relationships with macrobioclimate (MB) and bioclimates (BIO), and to the topographic and ecological gradients typical of coastal sandy areas. Results: Our bioclimate analysis served to define four MB and 11 BIO, each characterized by a particular type of potential natural vegetation. By floristically analysing the 522 vascular taxa detected, these were related to 16 phytogeographical elements. Syntaxonomically, the communities observed comprise 74 vegetation types, 70 of which are considered associations characterized by their diagnostic combination of species. These associations were classified into 24 alliances, 16 orders and 12 phytosociological classes. Two new classes, five new orders, 10 new alliances and 34 new associations are presented. Some syntaxa have been revised or validated. Conclusions: The coastal vegetation of the northern Pacific shows a distribution that is mainly linked to the four MB of the latitudinal gradient examined. Higher syntaxa are preferentially linked to a macrobioclimate, although some edaphic factors determine that some occupy two different but geographically close bioclimate zones. Regional factors, such as alkaline sands, induce the appearance of specialized vegetation types rich in endemic plants. Introduction This study of the beaches, dunes and their associated sandy plains from southern Alaska to Baja California is a survey the psammophilous plant communities of the NE Pacific using phytosociological methods. Despite some excellent regional works and good territorial floras, to date there is no general survey of the strand and dune vegetation of the North American Pacific coast. After an Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 1

Coastal vegetation of western North America Peinado, M. et al. initial period of pioneer studies clearly influenced by Cowles s (1899) theory of ecological succession on sand dunes, Cooper (1936) was the first to report the distribution of 53 representative coastal plants from SW Alaska to northern Baja California. Almost 40 yr later, Breckon & Barbour (1974), Macdonald & Barbour (1974) and Barbour et al. (1975) published the only known floristic syntheses of North America s Pacific beaches, but so far there have been no syntheses made of the plant communities of beaches and dunes in the Pacific coast of North America. Using the Braun-Blanquet method, 20 yr ago we started to study the coastal vegetation of the North American Pacific coast. So far, we have recorded 3259 relevés of coastal plant communities. These relevés constitute the basis of a series of phytosociological publications analyzing saltmarshes and mangroves (Delgadillo et al. 1992; Peinado et al. 1994b, 1995) and the coastal vegetation of Baja California (Peinado et al. 2008). The present study focuses on plant communities of the Pacific coast between the tip of the Baja California peninsula in Mexico, to the northernmost site at Cook Inlet, Alaska. The term coastal vegetation, as applied in this article, includes beach strand (not the beach itself but sparsely or densely vegetated areas behind the beach), foredunes, sand spits, and active to stable backdunes and sandsheets derived from quartz or alkaline sands. The vegetation of Mediterranean saltmarshes and tropical mangroves are outside the scope of this article, as these have been the subject of our above-mentioned studies. Estuarine wetlands and boreo-temperate saltmarshes are also excluded. Methods Study area Facing the Pacific Ocean, the study area extends across an airline distance of approximately 5000 km from the southern tip of Baja California (Loc. 1; for localities see Fig. S1) to Cook Inlet, Alaska (Loc. 279). In longitude, the area reaches its eastern limit at Bahía de Las Palmas (Loc. 2), while the westernmost site sampled was at Loc. 262, on Kodiak Island. The whole area forms part of the largest and highest of North American physiographic systems, the Pacific Border System (Brouillet & Whestone 1993), which is the backdrop for most of the ocean s shores (Fig. S2). Four zonobiomes (ZB) can be distinguished along this latitudinal band: ZB-VIII (cold-temperate or boreal), ZB-V (warm-temperate, maritime, humid), ZB-IV (winter rain and summer drought, arid-humid), and ZB-III or subtropical arid (Peinado et al. 1994a, 1997, 2007). From a phytogeographical standpoint, Dice (1943) included the boreal and the temperate climate zones within the Hudsonian (continental boreal), Sitkan (oceanic boreal) and Oregonian (temperate) provinces. The winter rain zone corresponds to the Californian Region, in which the provinces Northern California, Southern California, and Martirense have been defined. The tropical zone corresponds to the Xerophytic-Mexican region, which has been divided into two provinces: Baja Californian and Sanlucan (Fig. S2). For a more detailed phytogeographical classification see Peinado et al. (2008, 2009). Patterns of vegetation on Pacific dunes are best understood in the context of underlying geologic processes and dune morphology. Subsequent to Cooper s pioneering studies (1958, 1967), several authors have described dune geology and morphology for specific localities or regions. Using the information from these works, we provide a summary (Appendix S3) with the geological and geomorphological processes underlying coastal beaches and dunes, and a brief classification of beach and dune landscape as divided into major zones, each characterized by distinct plant habitats. See references in that table for additional and more extensive information on dune genesis. Selection of stands Prior to extensive fieldwork, some 300 potential strand localities were selected along the Pacific coast, from Cook Inlet, Alaska, to the southern tip of Baja California, including Kodiak and Vancouver islands. While the southern limit of the study area is marked by the extreme tip of the Baja California peninsula, its northern limit merely marks the end of the territory that we have explored. Localities were chosen to ensure an accessible strand and dune vegetation that was relatively well-preserved, and on the basis of satellite images, aerial photography, literature references, suggestions by other researchers, or our own knowledge of the coast. In subsequent fieldwork, we found that some of the localities selected were too disturbed by human activities. Finally, 279 localities were considered appropriate for detailed sampling (Fig. S1). These fall into several ecofloristic zones, ecoregions, and four zonobiomes (Appendix S4). Climate and potential natural vegetation (PNV) analyses Climate data from 226 coastal weather stations in the study area were compiled from the Canadian Weather Office (http://www.climate.weatheroffice.ec.gc.ca), NOAA (2002), and the Mexican National Weather Service (http://smn.cna. gob.mx). Several climate indices were calculated and every station was ascribed to one macrobioclimate (MB) and bioclimate (BIO). Four MB, grouping 11 BIO, can be distinguished across the study area: boreal, temperate, Mediterranean and tropical. For bioclimatic classification see Macías (2009) and Peinado et al. (2007, 2011a). Climate 2 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America diagrams are available at http://www2.uah.es/ambiente/ bioclimate/climograms.rar. Predicting vegetation type based on the structure or physiognomy of a community is the basic method of treating vegetation on a broad scale in relation to climate (Beard 1973). Accordingly, before the fieldwork, each weather station was initially assigned to a particular PNV, which is defined as the plant community that would become established if all successional sequences were completed without interference by man under the present climatic and edaphic conditions (Ricotta et al. 2002). In later fieldwork, this classification into vegetation types was checked. In this manner, the localities sampled were in turn assigned to a meteorological station and therefore to a BIO and PNV according to their geographical proximity. The bioclimatic classification and PNV for each weather station are summarized in Appendix S4. Bioclimates and PNV at association level along with climate data and indices for each station are available in Appendix S5. Field procedures This study is based on a set of 1730 relevés produced and analysed according to the Braun-Blanquet method (Westhoff & van der Maarel 1973; Braun-Blanquet 1979; Dierschke 1994). The localities were sampled from 1995 to 2009. Edaphic and geological data were previously obtained from bibliographic and cartographic sources. After reconnaissance of the entire locality, transects were defined along lines inland from the shoreline to the interior limit of the psammophilous vegetation. These transects represented a moisture or soil gradient (catena), such as from shoreline vegetation to dune ridges, or from the wet bottom of dune swales to the encircling drier dune slopes. Along each transect, zones were defined by changes in topography or vegetation. Significant vegetation changes included density, cover, and species composition. Our vegetation analysis started with the subjective choice of one or more stands at each transect zone following the approach termed subjective sampling without preconceived bias (Mueller-Dombois & Ellenberg 1974). Sampled stands were selected according to the homogeneity of physical features, structure of vegetation and dominance of species. Estimates of distance from the mean high tide, elevation, aspect, slope, position in the landscape, litter cover and parent material were recorded. Depending on the community type, plot sizes ranged from 1000 m 2, for forest stands, to 2 m 2, for some herbaceous communities. Coverabundance was estimated for vascular plants according to the six-point ordinal scale of Braun-Blanquet (1979). In total, 522 vascular taxa were recorded. Using several regional floras, plants were identified as far as possible in the field and specimens of doubtful identification collected for subsequent laboratory determination. These plants were deposited in the herbarium of the Universidad Autónoma de Baja California (BCMEX). Phytogeographical analysis The information available on the distribution range of each taxon recorded in the field was recompiled from the literature, mainly from the Flora of North America Editorial Committee (1993 2010), along with distribution maps obtained from the 2010 versions of three databases: Cal- Flora (http://www.calflora.org), E-Flora British Columbia (http://www.eflora.bc.ca) and USDA (http://plants.usda.- gov). According to their global distributions, each of the registered taxa was assigned to one of the 16 phytogeographical elements shown in Appendix S6. For more detailed information on these elements, see Peinado et al. (2007, 2009). Distribution ranges of the 522 recorded taxa is also shown in Appendix S6. In all the phytosociological tables species are scored with abbreviations (in superscript) to indicate they belong to one of those elements. Numerical analyses For the numerical analysis, Braun-Blanquet cover/abundance values were ordinally transformed: 1 was replaced by 1, 1 by 2, 2 by 4, 3 by 6, 4 by 7 and 5 by 8. Analyses were performed using SPSS version 13.0 software (IBM, Somers, NY, US), starting with a data set (1730 relevés) that included each plant species recorded in the relevés (522), together with its respective transformed cover/ abundance value. Classification was based on squared Euclidean distances and the Average Linkage Clustering method (ALC; Wildi 2010). A stepwise procedure of successive approximations through ALC and rearrangement of synoptic tables was used to classify our field data. A total of nine ALC were performed in two stages (for dendrograms see Appendix S7). In an initial ALC using the starting data set, 1691 relevés were included in 50 clusters or relevé groups (RG). The remaining 39 relevés, individually or in pairs, appeared distributed throughout the dendrogram. Combining clustering results with tabular rearrangement, literature survey, and field notes, 21 RG were directly related to already described Baja Californian associations (Table 1: associations 26 to 41, 43, 46 and 48 to 55). Depending on their floristic, phytogeographical and ecological affinities, eight new data sets were built with the 29 remaining RG (Appendix S8). Of the 39 scattered relevés, 12 were included in one of these new data sets according to their floristic affinities, and 27 were removed because they did not correspond to true psammophilous vegetation types. As a new step, these new eight data sets were independently subjected to a further ALC, with the Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 3

Coastal vegetation of western North America Peinado, M. et al. Table 1. Complete syntaxonomical scheme and brief diagnoses of the new syntaxa. Complete diagnoses are done with the descriptions made along the main text and in App. 1. For the diagnostic species of each syntaxon see Appendices S10 and S11. Phytosociological tables for new associations in Appendix S12. Syntaxa marked with asterisk ( ) are typified in App. 1. (I) Honckenyo Elymetea arenarii Tx. 1966. (I.1) Honckenyo majoris Elymetalia mollis Ohba, Miyawaki et Tx. 1973. (I.1.1) Senecioni pseudoarnicae Leymion mollis Ohba, Miyawaki et Tx. ex Peinado et al. 2011. (1) Mertensio maritimae Honckenyetum majoris Peinado et al. 2011, boreal tide-mark association, occurring at the leading edge of the vegetation, periodically inundated by ocean waves and exposed to salt spray on well-drained gravelly and stony beaches, and on river and creek mouths prone to freshwater floods (Table S12.1). (2) Senecioni pseudoarnicae Leymetum mollis Peinado et al. 2011, boreal beach and foredune meadows periodically influenced by salt spray (Table S12.2). (3) Lathyro palustris Leymetum mollis Peinado et al. 2011, boreal foredune wet meadows, that substitutes association 2 inland, in ecotones with mesic graminoid meadows dominated by Calamagrostis canadensis (Table S12.3). (4) Lathyro maritimi Leymetum mollis Peinado et al. 2011, temperate beach and foredune meadows (Table S12.4). (5) Leymo mollis Caricetum macrocephalae Peinado et al. 2011, temperate beach meadow characterized by the dominance of Carex macrocephala, a perennial sedge that spreads rhizomatously across the upper beach and among the jumble of driftwood at the furthest reach of winter storm tides and waves (Table S12.5). (6) Leymo mollis Ammophiletum arenariae Peinado et al. 2011, foredune-builder naturalized vegetation on temperate sandy beaches (Table S12.6). (II) Cakiletea maritimae Tx. et Prsg. ex Br.-Bl. et Tx. 1952. (II.2) Cakiletalia edentulae (Tx. 1950) Thannheiser 1981. (II.2.2) Cakilion edentulae Thannheiser 1981. (7) Cakiletum maritimo edentulae Peinado et al. 2011, temperate tide-mark association, occurring at the leading edge of the vegetation, periodically inundated by ocean waves and exposed to salt spray (Table S12.7). (III) Ambrosietea chamissonis Kohler 1970 (III.3). Abronietalia latifoliae Knapp ex Peinado et al. 2011. Rhizomatous upper-beach and foredune vegetation of the Northern Californian phytogeograhical province (from Morro Bay northwards). (III.3.3) Ambrosion chamissonis Ohba, Miyawaki et Tx. ex Biondi et Casavecchia 2001. (8) Abronietum latifoliae Peinado et al. 2011, association dominated by perennial herbs forming shadow dunes when colonizes the upper beach, and building discontinuous hummocks when thriving on the seaward edge of dunes. (8b) Ambrosietosum chamissonis Peinado et al. 2011, on the leeward side of the hummocks. (8c) leymetosum mollis Biondi et Casavecchia stat. nov., on wind-sheltered, rear mounds. (8d). In some sites leeward slopes are colonized by pure communities of Ambrosia chamissonis (Table S12.8). (9) Lupino variicoloris Ammophiletum australis Biondi & Casavecchia 2001. (III.4) Abronietalia maritimae Knapp ex Peinado et al. 2008. (III.4.4) Atriplici leucophyllae Abronion maritimae Knapp ex Peinado et al. 2008. Rhizomatous upper-beach and foredune vegetation of the Southern Californian and Martirense phytogeograhical provinces (from Morro Bay southwards). Ten. Ambrosio chamissonis Abronietum maritimae Peinado et al. 2011, Thermo-mediterranean semi-arid association dominated by perennial herbs forming shadow dunes when colonizes the upper beach, and building conical hillocks on the seaward edge of dunes. (10b) Ambrosietosum chamissonis Peinado et al. 2011, on the leeward side of the hillocks. (10c) In some sites leeward slopes are colonized by pure communities of Ambrosia chamissonis. (10d) atriplicetosum leucophyllae Biondi et Casavecchia stat. nov., forming shadow dunes and nebkhas in the open area of the strand. (10e) Pure communities of Atriplex leucophylla growing rather close to the ocean, often near strandlines, along with Abronia seedlings and Cakile singly plants, are assigned to the Atriplex leucophylla community (Table S12.9). (10f) Variant of Leymus mollis, on the leeward side of the hillocks at Morro Bay. (11) Abronietum maritimae Peinado et al. 2008. 12. Isocomo menziesii Ambrosietum chamissonis Peinado et al. 2008 (III.5) Ambrosio chamissonis Eriogonetalia latifolii Peinado et al. 2011. Oceanic dune-scrubs: mesophanerophytic, nanophanerophytic, chamaephytic and suffrutescent vegetation growing on aeolian sands that constitutes the psammophilous edaphoclimax on half-stabilized and stabilized backdunes of the Californian phytogeographical region. Communities of this order replaces inland to the foredune vegetation of Abronietalia maritimae and Abronietalia latifoliae. (III.5.5) Artemision pycnocephalae Peinado et al. 2011, dune mats living on ridges, half stabilized mid dunes and in prograding shores near river mouths, from Monterey northward. (13) Ambrosio chamissonis Artemisietum pycnocephalae Peinado et al. 2011, flourishes on ecotones between the herbaceous vegetation of foredunes and the typical dune scrubs on backdunes (Table S12.10). (14) Eriogono parvifolii Artemisietum pycnocephalae Peinado et al. 2011, on dune ridges and leeward slopes in the Monterey Bay dune complex (Table S12.11). (15) Heterotheco bolanderi -Artemisietum pycnocephalae Peinado et al. 2011, on dune ridges and leeward slopes in the Ten Mile River dune complex (Table S12.12). (16) Polygono paronychiae Artemisietum pycnocephalae Biondi & Casavecchia 2001. 17. Poetum douglasii Peinado et al. 2011, pioneer dry meadow on blowouts and other inland sites with reworked sand (Table S12.13). (III.5.6) Lupinion arboreo chamissonis Peinado et al. 2011, open dune scrubs on windward, more exposed slopes, usually on re-worked sands. (18) Agoserido eastwoodieae Lupinetum chamissonis Peinado et al. 2011, lupine scrubs from the Northern Californian province (Table S12.14). (19) Senecioni blochmaniae Lupinetum chamissonis Peinado et al. 2011, lupine scrubs from the Southern Californian province. Variant of Ericameria ericoides on ecotones with Corethrogyno californicae Ericamerietum ericoidis (Table S12.15). (20) Achilleo arenicolae Lupinetum arborei Peinado et al. 2011, nitrophilous dune shrubs on inland, disturbed, sites from the Northern Californian province. (20b) Eriophylletosum staechadifolii Peinado et al. 2011, thriving on exposed slopes nearest the ocean (Table S12.16). (III.5.7) Ericamerion ericoidis Peinado et al. 2011, dense dune scrubs on stabilized lee slopes. (21) Baccharido pilularis Ericamerietum ericoidis Peinado et al. 2011, dune scrubs living on entirely stabilized hind dunes and lee slopes from the Northern Californian province (from Monterey Bay northward) (Table S12.17). (22) Corethrogyno californicae Ericamerietum ericoidis Peinado et al. 2011, dune scrubs living on entirely stabilized hind dunes and lee slopes from the Southern Californian province (Table S12.18). (23) Dudleyo farinosae Ericamerietum ericoidis Peinado et al. 2011, dune scrubs living on entirely stabilized hind dunes and lee slopes. Endemic to Monterey Bay dunes (Table S12.19). (III.5.8) Erigeronto glauci Eriophyllion staechadifolii Peinado et al. 2011, dense dune scrubs on sea bluffs. (24) Eriogono parvifolii Eriophylletum staechadifolii Peinado et al. 2011, coastal scrub living on the ocean bluffs south of Monterey Bay (Table S12.20). (25) Erigeronto glauci Eriophylletum staechadifolii Peinado et al. 2011, coastal scrub living on the ocean bluffs north of Monterey Bay (Table S12.21). 4 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America Table 1. Continued (IV) Atriplici julaceae Frankenietea palmeri Peinado et al. 2008 (IV.6) Frankenietalia palmeri Peinado et al. 2008 (IV.6.9) Atriplici julaceae Frankenion palmeri Peinado et al. 2008. 26. Atriplici linearis Frankenietum palmeri Peinado et al. 2008. 27. Atriplici julaceae Frankenietum palmeri Peinado et al. 2008. 28. Euphorbio miserae Lycietum californici Peinado et al. 2008. 29. Dudleyo cultratae Lycietum californici Peinado et al. 2008. (IV) (7) Camissonio crassifoliae Isocometalia menziesii Peinado et al. 2008. (IV.7.10) Heliantho nivei Isocomion menziesii Peinado et al. 2008. 30. Loto bryanthii Isocometum menziesii Peinado et al. 2008. 31. Heliantho nivei Isocometum vernonioidis Peinado et al. 2008. 32. Camissonio crassifoliae Helianthetum nivei Peinado et al. 2008. 33. Heliantho nivei Astragaletum anemophili Peinado et al. 2008. (IV.7.11) Encelion ventori Peinado et al. 2008. 34. Camissonio crassifoliae Encelietum ventori Peinado et al. 2008. 35. Sphaeralceo fulvae Encelietum ventori Peinado et al. 2008. (IV.7.12) Lycion richii Peinado et al. 2008. 36. Lycietum brevipedis Peinado et al. 2005. (37) Ephedro californicae Lycietum richii (Peinado et al. 2005) Peinado et al. 2008. (V) Achyronichio cooperi Abronietea villosae Peinado et al. 2008. (V.8) Nicolletio trifidae Verbenetalia bajacalifornicae Peinado et al. 2008. (V.8.13) Chaenactido lacerae Dyssodion anthemidifoliae Peinado et al. 2008. 38. Plantagini ovatae Chaenactidetum lacerae Peinado et al. 2008. (VI) Stellarietea mediae Tx., Lohm. et Prs. ex von Rochow 1951. (VI.9) Chenopodietalia muralis Br.-Bl. in Br.-Bl., Gajewski, Wraber et Walas 1936. (VI.9.14) Mesembryanthemion crystallini Rivas-Martínez et al. 1993. (39) Amblyopappo pusilli Mesembryanthemetum crystallini Peinado et al. 2008. (VII) Prosopido torreyanae Fouquierietea splendentis Rivas-Martínez 1997 (VII.10) Parkinsonietalia florido-microphyllae Rivas-Martínez 1997. (VII.10.15) Prosopido Carnegeion gigantei Rivas-Martínez 1997. (40) Errazurizio megacarpae Ephedretum trifurcae Peinado et al. 2006. (41) Pachycereo schottii Prosopidetum torreyanae Peinado et al. 2006. (VII.11) Opuntio chollae Stenoceretalia gummosae Peinado et al. 2011, drought deciduous and succulent scrubs, and microphyll woodlands living on the tropical (thermotropical and mesotropical belts with hyperarid to arid ombroclimates) deserts of the Baja California peninsula (Baja Californian and Sanlucan phytogeographical provinces). Communities of this order also penetrate into the coastal infra-mediterranean desert of El Vizcaíno, in the southernmost corner of the Martirense province. (VII.11.16) Ferocacto towsendiani Fouquierion diguetii Peinado et al. 2011, drought deciduous and succulent scrubs, and microphyll woodlands living on the thermotropical deserts of the Baja California peninsula (Baja Californian and Sanlucan phytogeographical provinces). (42) Jatropho cordate Cyrtocarpetum edulis Peinado et al. 2011, sarcocaulescent thornscrubs dominated by columnar cacti and deciduous sarcocaulescent small trees growing on arenosols in the sandy inland plains immediately behind the Sanlucan dune systems (Table S12.22). (43) Yucco validae Fouquierietum diguetii Peinado et al. 1995. (VII.11.17) Lycio congesti Maytenion phyllanthoidis Peinado et al. 2011, dense thickets dominated by Maytenus phyllanthoides accompanied with drought deciduous and succulent scrubs living on sea-exposed slopes with an inmediate influence of salt spray. (44) Cyrtocarpo edulis Maytenetum phyllanthoidis Peinado et al. 2011, M. phyllanthoides dense thickets thriving on the leeward slopes of the Sanlucan backdunes (Table S12.24). (45) Lycio congesti Maytenetum phyllanthoidis Peinado et al. 2011, Maytenetum phyllanthoides dense and flagged thickets thriving on the windward slopes of the Sanlucan backdunes (Table S12.23). (VIII) Allenrolfeetea occidentalis Peinado et al. 2008. (VIII.12) Allenrolfeetalia occidentalis Peinado et al. 2008. (VIII.12.18) Allenrolfeion occidentalis Peinado et al. 2008. 46. Suaedo taxifoliae -Allenrolfeetum occidentalis Peinado et al. 2006. (IX) Euphorbio leucophyllae Sporoboletea virginici Peinado et al. 2008. (IX.13) Sporobolo virginici Jouveetalia pilosae Peinado et al. 2008 (IX.13.19) Palafoxio linearis Abronion maritimae Peinado et al. 2008. Hillock-builder associations thriving in the northern part of the Xerophytic- Mexican region, from central Baja California to Sinaloa in continental Mexico. (47) Sarcostemmato arenarii Astragaletum magdalenae Peinado et al. 2008. 48. Atriplicetum magdalenae Peinado et al. 2008. 49. Palafoxio linearis Daleetum tinctoriae Peinado et al. 2008. 50. Daleo maritimae Jouveetum pilosae Peinado et al. 2008. (IX.13.20) Oenothero talassaphilae-jouveion pilosae Peinado et al. 2008. Beach and foredune grasslands endemic to southern Baja California (Sanlucan province and neighboring areas). (51) Euphorbio leucophyllae Drymarietum crassifoliae Peinado et al. 2008. (52) Euphorbio leucophyllae Jouveetum pilosae Peinado et al. 2008. (53) Sporobolo virginici Ipomoeetum brassiliensis Peinado et al. 2008. 54. Ipomoeo imperati Jouveetum pilosae Peinado et al. 2008. (55a) Daleo anthonyi Jouveetum pilosae Peinado et al. 2008. (55b) Sporobolus virginicus community. (X) Tsugetea mertensiano heterophyllae Rivas-Martínez et al. 1999. (X.14) Tsugetalia mertensiano heterophyllae Rivas-Martínez et al. 1999. (X.14.21) Pinion contortae Rivas-Martínez et al. 1999. (56) Carici obnuptae-pinetum contortae Peinado et al. 2011b. (57) Arctostaphylo uva-ursi-pinetum contortae Peinado et al. 2011a, b. (58) Lonicero ledebourii Pinetum contortae Rivas-Martínez et al. 1999. (59) Rhododendro macrophylli Pinetum contortae Peinado et al. 2011b. (60) Morello californicae Piceetum sitchensis Peinado et al. 2011b. (XI) Salicetea lasiandro exiguae Peinado et al. 2011a, b, riparian willow thickets. Open to dense broadleaved deciduous tickets typically o 10 m in height, dominated by spindly shrubs, many with clonal growth by root sprouting, often accompanied by multi-stemmed small willow trees (2 12 m), and scattered taller willow trees (Salix gooddingii, Salix laevigata). Riparian willows are tolerant to frequent flooding and sustained inundation, occurring on streamside areas, river islands and oxbox lakes, lakeshores and wet meadows, often standing in quiet, shallow river backwaters. Riparian willows also replace riparian cottonwood, sycamore and red alder forests on exposed gravel bars. On coastal dunes, willow thickets occur on waterlogged swales. Currently, the class includes two new orders: Morello californicae Salicetalia,andSalicetalia delnortensisbreweri Peinado et al. 2011a, b, which encloses riparian thickets occupying ultramafic streambeds, creeks and canyons in lowlands throughout the Northern Californian province (Coast Ranges, Klamath-Siskiyou Mountains). (XI.15) Morello californicae Salicetalia Peinado et al. 2011a, b. (XI.15.22) Morello californicae Salicion Peinado et al. 2011a, b, Mediterranean and temperate willow shrubs on dune swales and deflation plains of the Pacific coast of western North America. (61) Morello californicae Salicetum bigelovii Peinado et al. 2011a, b, Mediterranean willow shrubs (Table S12.25). (62) Lonicero ledebourii Salicetum hookerianae Peinado et al. 2011a, b, northern Mediterranean and temperate submediterranean willow shrubs (Table S12.26). (63) Lonicero involucratae Salicetum hookerianae Peinado et al. 2011a, b, temperate willow shrubs (Table S12.27). Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 5

Coastal vegetation of western North America Peinado, M. et al. Table 1. Continued (XII) Juncetea breweri Peinado et al. 2011a, b. (XII.16) Juncetalia breweri Peinado et al. 2011a, b, herbaceous, usually rhizomatous, vegetation on oligotrophic habitats. (XII. 16.23) Argentino egedii Caricion obnuptae Peinado et al. 2011a, b, wet sedge and rush meadows thriving on periodically waterlogged habitats. (64) Argentino egedii Caricetum obnuptae Peinado et al. 2011a, b, Mediterranean sedge meadows, on sites where water persists 4 to 6 months (Table S12.28). (65) Veronico scutellatae Caricetum obnuptae Peinado et al. 2011, temperate sedge meadows, on sites where water persists four to 6 months (Table S12.29). (66) Juncetum breweri falcati Peinado et al. 2011, Mediterranean rush meadows on wet habitats where water persists on the surface for three or 4 months in winter (Table S12.30). (67) Juncetum breweri-sitchensis Peinado et al. 2011, vicariant of the previous association in temperate areas (Table S12.31). (68) Junco breweri Caricetum pansae Peinado et al. 2011, sedge meadows on habitats where water persists on the surface for 1 or 2 months in winter (Table S12.32). (XII.16.24) Lupino littoralis Polygonion paronychiae Peinado et al. 2011, dry meadows thriving on dune crests and on the driest areas of the deflations plains. (69) Poo macranthae Lupinetum littoralis Peinado et al. 2011, temperate open dry meadows (Table S12.33). (70) Polygono paronychiae Tanacetetum camphorati Peinado et al. 2011, dry meadows in prograding shores. (70a), Tanacetetosum camphorati Peinado et al. 2011, temperate. (70b) Camissonietosum cheiranthifoliae Peinado et al. 2011, Mediterranean (Table S12.34). aim of defining floristic groups in more detail. For each of the resultant RG, new synoptic tables were prepared by scoring species for presence percentages (Appendix S9). Syntaxonomical analysis The syntaxonomical analysis was performed in two successive steps. In the first step, the synoptic tables obtained in the numerical analysis served for a further tabular rearrangement using presence percentages and mean cover expressed as cover/abundance classes of Braun- Blanquet (1979). Comparisons of the frequencies in the synoptic tables in conjunction with expert judgement resulted in a vegetation typology based on similarities in presence/absence and cover/abundance values of the species. As a result, preliminary floristic groups were obtained. These were interpreted as vegetation types and used to generate a new presence table, which served to establish relationships among them and form the basis for a preliminary syntaxonomical treatment. To test the strength of this preliminary classification, in the second step diagnostic species for preliminary syntaxa were determined by fidelity calculations using the phi-coefficient of association (Chytrý et al. 2002a, b). For the statistical analysis of diagnostic species rare species (presence o 10% in any association) were eliminated. Thus, the final data set included 1703 relevés and 392 species. Calculations were done using the JUICE 7.0 program (http://www.sci.muni.cz/botany/juice.htm). The hierarchical syntaxonomical vegetation classification system according to the Braun-Blanquet method consists of the units association, alliance, order, and class, which were delimited by their diagnostic species. The term communities, as we consider here, refers to recognizable vegetation units, but without diagnostic species; thus, they are not part of the formal hierarchy. The descriptions of the new phytosociological classes and, in general, of all the higher syntaxa proposed in this article, follow the concept of Pignatti et al. (1995). Nomenclature and typification of syntaxa is in agreement with the International Code of Phytosociological Nomenclature (ICPN; Weber et al. 2000). Results and Discussion As a consequence of table rearrangements supported by determination of diagnostic species, the initial 50 RG were finally classified into 70 associations and four communities. Main physiognomic, edaphic and ecological features of the recognized syntaxa are summarized in Table 1. Complete syntaxonomical tables in Appendix S10 show the floristic relations of associations with higher syntaxa. Summary Table 2 and Appendix S11 show diagnostic species and phi-values for higher syntaxa and associations, respectively. Nomenclatural type relevés for each association and diagnoses for higher syntaxa are shown in App. 1. The complete tables for new associations may be found in Appendix S12, and a dichotomous key to associations based on floristic, phytogeographical and ecological features is provided in Appendix S13. Boreal and temperate beach and foredune grasslands (Table S10.1) On beaches and foredunes, four major types of vegetation occur: (1) boreal, temperate and northern Mediterranean grasslands with Leymus mollis subsp. mollis (hereafter L. mollis); (2) tidemark pioneer communities dominated by Cakile species; (3) Mediterranean hillock-builder dune mats, dominated by perennial forbs, and (4) tropical grasslands with Euphorbia leucophylla and Jouvea pilosa. In our study area, meadows dominated or co-dominated by L. mollis occur on Mediterranean, temperate and boreal MB, but the floristic assemblage of those communities differs such that they may be included within two phytosociological classes: Honckenyo Elymetea arenarii and Ambrosietea chamissonis. Boreal and temperate L. mollis meadows are assigned to the amphi-pacific alliance Senecioni pseudoarnicae Leymion 6 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America Table 2. Synoptic table of relevés (N = 1703) with diagnostic taxa. Syntaxa codes as in Table 1. For each syntaxon, the most faithful species (according to their phi-coefficient) are listed in descending order. f =Phicoefficient ( 1000 and rounded). Np = number of relevés belonging to the syntaxon. Only classes, orders and alliances and species with phivalues 4 0.3 are shown. For additional diagnostic species and diagnostic species of associations see Appendix S11. Class I. Order 1. Alliance 1. Np: 201 f Leymus mollis subsp. mollis NAE 760 Lathyrus japonicus v. maritimus NAE 721 Honckenya peploides subsp. major NAE 416 Senecio pseudoarnica NAE 390 Achillea millefolium v. borealis HOL 376 Class II. Order 2. Alliance 2. Np:31 Cakile edentula subsp. edentula v. edentula INT 827 Cakile maritima subsp. maritima INT 401 Atriplex patula COS 316 Class III. Np: 523 Ambrosia chamissonis NAS 476 Ericameria ericoides MED 416 Abronia latifolia PAC 422 Artemisia pycnocephala MED 410 Eriogonum latifolium MED 385 Order 3. Alliance 3. Np: 119 Abronia latifolia PAC 476 Ammophila arenaria INT 413 Order 4. Alliance 4. Np: 88 Abronia maritima MAD 716 Cakile maritima subsp. maritima INT 342 Malacothrix incana MED 337 Atriplex leucophylla MED 326 Order 5. Np: 315 Ericameria ericoides MED 410 Eriophyllum staechadifolium MED 338 Artemisia pycnocephala MED 366 Eriogonum latifolium MED 357 Achillea millefolium v. arenicola MED 326 Alliance 5. Np: 106 Artemisia pycnocephala MED 764 Ambrosia chamissonis NAS 509 Eriophyllum staechadifolium MED 338 Polygonum paronychia PAC 304 Alliance 6. Np:95 Lupinus chamissonis MED 434 Lupinus arboreus PAC 364 Phacelia ramosissima v. austrolitoralis MED 328 Senecio blochmaniae MED 306 Alliance 7. Np:71 Ericameria ericoides MED 683 Croton californicus MAD 359 Conicosia pugioniformis INT 308 Alliance 8. Np:44 Eriophyllum staechadifolium MED 597 Dudleya farinosa MED 452 Erigeron glaucus MED 489 Angelica hendersonii PAC 449 Rubus ursinus subsp. ursinus WES 374 Class IV. Np: 224 Atriplex julacea BAJ 719 Table 2. Continued Frankenia palmeri MAD 643 Euphorbia misera MAD 455 Lycium richii MAD 441 Lycium californicum MAD 414 Order 6. Alliance 9. Np: 101 Frankenia palmeri MAD 715 Euphorbia misera MAD 530 Lycium californicum MAD 513 Order 7. Np: 123 Isocoma menziesii v. menziesii MAD 429 Helianthus niveus subsp. niveus MAD 406 Lycium richii MAD 378 Encelia ventorum BAJ 358 Camissonia crassifolia BAJ 314 Alliance 10. Np: 55 Helianthus niveus subsp. niveus MAD 632 Isocoma menziesii v. menziesii MAD 543 Astragalus anemophilus BAJ 382 Lotus bryantii BAJ 310 Alliance 11. Np: 36 Encelia ventorum BAJ 941 Camissonia crassifolia BAJ 765 Sphaeralcea fulva BAJ 445 Haplopappus sonorensis SON 382 Viguiera deltoidea v. chenopodina BAJ 352 Alliance 12. Np: 32 Lycium richii MAD 781 Atriplex julacea BAJ 348 Class V. Order 8. Alliance 13. Np: 26 Plantago ovata MAD 960 Chaenactis lacera BAJ 883 Phaseolus acutifolius v. latifolius NEO 853 Dyssodia anthemidifolia BAJ 782 Abronia umbellata subsp. umbellata MAD 706 Class VI. Order 9. Alliance 14. Np: 14 Amblyopappus pusillus NAS 1000 Mesembryanthemum nodiflorum INT 925 Lepidium nitidum MAD 706 Bromus rubens INT 596 Mesembryanthemum crystallinum INT 574 Class VII. Np: 95 Maytenus phyllanthoides NEO 689 Pachycereus pringlei SON 643 Lycium fremontii v. congestum SON 618 Cyrtocarpa edulis BAJ 575 Larrea tridentata v. tridentata NEO 575 Order 10. Alliance 15. Np: 21 Ambrosia dumosa SON 813 Prosopis glandulosa v. torreyana MAD 744 Larrea tridentata v. tridentata NEO 678 Ephedra trifurca SON 530 Hilaria rigida SON 530 Order 11. Np: 74 Maytenus phyllanthoides NEO 518 Lycium fremontii v. congestum SON 428 Cyrtocarpa edulis BAJ 383 Stenocereus gummosus SON 331 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 7

Coastal vegetation of western North America Peinado, M. et al. Table 2. Continued Alliance 16. Np: 27 Yucca valida BAJ 701 Fouquieria diguetii SON 701 Jatropha cinerea NEO 621 Larrea tridentata v. tridentata NEO 593 Opuntia ciribe BAJ 565 Alliance 17. Np: 47 Maytenus phyllanthoides NEO 1000 Lycium fremontii v. congestum SON 782 Jatropha cuneata SON 359 Class VIII. Order 12. Alliance 18. Np: 13 Allenrolfea occidentalis MAD 785 Suaeda taxifolia MAD 649 Class IX. Order 13. Np: 158 Euphorbia leucophylla SON 686 Oenothera drummondii v. thalassaphila BAJ 612 Jouvea pilosa NEO 553 Ipomoea pes-caprae subsp. brasiliensis NEO 400 Proboscidea althaefolia SON 406 Alliance 19. Np: 41 Abronia maritima MAD 701 Atriplex magdalenae BAJ 589 Chamaesyce micromera NEO 548 Dalea tinctoria v. tinctoria BAJ 535 Alliance 20. Np: 117 Oenothera drummondii v. thalassaphila BAJ 477 Jouvea pilosa NEO 346 Euphorbia leucophylla SON 423 Proboscidea althaefolia SON 298 Drymaria holosteoides v. crassifolia BAJ 250 Class X. Order 14. Alliance 21. Np: 160 Gaultheria shallon PAC 844 Vaccinium ovatum PAC 820 Pinus contorta v. contorta PAC 806 Picea sitchensis PAC 752 Morella californica PAC 634 Class XI. Order 15. Alliance 22. Np: 81 Salix lasiolepis v. bigelovii MED 629 Salix hookeriana PAC 542 Scrophularia oregana TEM 361 Lonicera involucrata v. ledebourii MED 364 Class XII. Order 16. Np: 177 Juncus falcatus v. sitchensis TEM 535 Carex obnupta PAC 487 Veronica scutellata NOA 478 Carex pansa PAC 467 Juncus breweri TEM 593 Alliance 23. Np: 135 Carex obnupta PAC 555 Juncus falcatus v. sitchensis TEM 384 Argentina egedii subsp. egedii NAE 384 Carex pansa PAC 355 Veronica scutellata NOA 321 Alliance 24. Np: 42 Tanacetum camphoratum WES 675 Polygonum paronychia PAC 856 Poa macrantha PAC 651 Ammophila arenaria INT 483 mollis (Honckenyo majoris Elymetalia mollis), within the circumboreal class Honckenyo Elymetea arenarii. Several amphi-beringian taxa are present in both Japanese and North American associations (see NAE taxa in Table S10.1). Such relationships result from the ancient intercontinental connection between eastern Asia and western North America across the Bering bridge from the Late Cretaceous to the Pleistocene (Peinado et al. 2009). This alliance encompasses six associations. Mertensio maritimae Honckenyetum majoris is an open and species-poor tidemark pioneer association common in the boreal part of the study area flourishing between the mean high and the extreme high tidal level; because it is periodically inundated by ocean waves, drift belts of organic matter are often present covering plants after high tides. Beach meadows with a similar ecology and dominated by M. maritima and/or H. peploides have been described from northern Europe, Greenland, eastern North America and Canada (Lepping & Daniëls 2007) to continental Alaska (Boggs 2000), the Aleutian Islands (Talbot et al. 2010), and eastern Asia (Ohba et al. 1973). Griggs (1936) emphasized the role of M. maritima and H. peploides subsp. major as pioneer plants colonizing the newly formed beaches after the eruption of the Katmai, Alaska. The nurse-plant role of H. peploides in the very first stages of primary succession on the upper beach facilitates the recruitment of L. mollis by trapping seeds and improving seed germination and seedling emergence (Gagné & Houle 2001). Communities dominated by an upper field layer (height rangesfrom0.5to1.5m)ofl. mollis, inassociationwith many forbs, occur as tall, dense, usually linear stands oriented parallel to the shore on gravelly, sandy or finer substrates at the storm line. In boreal areas, we recognize two associations: dry beach grasslands (Senecioni pseudoarnicae Leymetum mollis), and mesic beach grasslands (Lathyro palustris Leymetum mollis). The former is the second colonizer of foredunes, flourishing on well-drained sites on level to steep slopes where it forms an inland zone as the frequency of tidal flooding decreases, immediately behind Mertensio maritimae Honckenyetum majoris, the initial pioneer at or above beach storm lines. Lathyro palustris Leymetum mollis substitutes Senecioni pseudoarnicae Leymetum mollis inland, in ecotones with mesic graminoid meadows dominated by Calamagrostis canadensis, which are transitional to coastal and estuarine wetlands. Since the first descriptions by Hanson (1951), the L. mollis foredune meadow and H. peploides beach meadow are referred to different Elymus mollis vegetation types, which are summarized by Viereck et al. (1992: 38). For additional references see Boggs (2000) and Talbot et al. (2010). In temperate areas there are two associations dominated by native species: Leymo mollis Caricetum macrocephalae, a beach meadow growing among driftwoods, and Lathyro maritimi Leymetum mollis, a foredune 8 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America grassland, which now appears in small populations scattered across temperate coasts because its original area has been vastly reduced by the spread of marram grass, Ammophila arenaria, the most pervasive dune invader of those coasts. Continuous foredune ridges similar to those on the white and grey coastal sand dunes of Europe did not originally exist on the Pacific coast of North America. Before the introduction of A. arenaria, the foredune was generally a series of closely spaced mounds formed by native species, with L. mollis as dominant or co-dominant (Cooper 1967). The continuous and steep foredunes of northern California, Oregon and Washington are a recent phenomenon developed mostly since 1868, when A. arenaria was first introduced to California as a first sand binder to stabilize the dunes in San Francisco (Lamb 1898), and for use in dune control planting on the Oregon coast (Wiedemann & Pickart 2004). On the temperate coasts, A. arenaria has built a steep, continuous foredune in the immediate shore area, where sand deposition by wind is greatest, replacing the original low, hummocky native foredunes and drastically reducing the amount of sand moving inland off the beach. Along most of those coasts, the foredune has attained heights of 10 m and a base width of over 100 m. These temperate foredune grasslands are here assigned to Leymo mollis Ammophiletum arenariae, in which Ammophila occurs at cover values of up to 99% with few associated species that are the remains of the older Lathyro maritimi Leymetum mollis. Temperate and Mediterranean ephemeral tidemark vegetation (Table S10.2) The annual vegetation of drift lines (Cakiletea maritimae)is widely distributed across the study area, but only under Mediterranean and temperate MB. It is absent under boreal MB areas because of the short vegetation season and the temporary habitat, which is frequently disturbed by waves and drift ice. Such conditions do not allow therophytes to complete their life cycle in circumboreal areas (Lepping & Daniëls 2007). Cakiletea maritimae is also absent from tropical areas around the world (Doing 1985). In our study area, the tropical tidemark vegetation is dominated by perennial species, which have been assigned to associations belonging to Euphorbio leucophyllae-sporoboletea virginici (Peinado et al. 2008). Although in our study area there are no native Cakile species (Rodman 2010), annual tidemarks are dominated by Cakile edentula subsp. edentula var. edentula (hereafter C. edentula), and C. maritima subsp. maritima (hereafter C. maritima) on temperate beaches, and only by C. maritima on Mediterranean shores. Now, C. maritima, introduced from Europe, is replacing its introduced precursor C. edentula (native to eastern North America) in central and southern California as a result of its ability to survive into a second or third reproductive season (Boyd & Barbour 1993), thus explaining the observed disappearance of C. edentula along the Mediterranean coast of California (Barbour & Rodman 1970). On temperate coasts, species coexistence has been attributed to a strictly annual habit forced upon both species by more severe winter conditions (Pickart & Barbour 2007). Couch (1914), Ramaley (1918), Purer (1936) and Pierce and Pool (1938) provided historical accounts of Californian strand vegetation. None of these authors documented the presence of annual, tidemark communities, but they did mention four perennial species that colonized the upper beach immediately above the debris line: Abronia latifolia, Abronia maritima, Atriplex leucophylla, and Camissonia cheiranthifolia. Only A. leucophylla and C. cheiranthifolia spread throughout the Mediterranean MB zone, and both are the most aggressive of foredune builders in their advance toward the sea, building shadow, ephemeral mounds just above the mean high-water spring tides. Perennial communities of A. leucophylla and C. cheiranthifolia have been assigned here to Ambrosietea chamissonis. Currently, on some Californian beaches where the vegetation has not been completely destroyed by human activities such as trampling, beach cleaning and recreational use, the vegetation colonizing embryo dunes is dominated by communities of C. maritima covering scattered individuals of A. leucophylla half buried by the sand (see Atriplex leucophylla community on Table S12.9). Historical data on the original composition of tidemark communities in temperate areas are not available, but the presence among driftwoods of species such as Calystegia soldanella, Carex macrocephala, Glehnia leiocarpa or L. mollis, suggests they could be the native dominants of the temperate tidemark vegetation. Communities dominated by both sea-rockets now flourish in the temperate zone and have been included in the new association Cakiletum maritimo-edentulae. Mediterranean beach and foredune vegetation (Tables S10.1, S10.2 and S10.3) While the fruticose dune vegetation on alkaline sands from Baja California was included within the class Atriplici Frankenietea palmeri (Peinado et al. 2008), the beach and dune perennial vegetation on quartz sands that occurs along the Mediterranean climate coasts of both Americas is here assigned to the class Ambrosietea chamissonis. The amphi-american distribution of this class on Chilean and Californian shores was mentioned in its original description (Kohler 1970: 96) and later confirmed (Kohler 1975; Eskuche 1992). Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 9

Coastal vegetation of western North America Peinado, M. et al. The class is supported by plants that thrive on coastal dunes in Chile and in the Californian phytogeographical region (see NAS taxa in Appendix S5). These disjunctions have been related to hydrochloric seed dispersal by ocean streams (Kohler 1975; Ramírez & Romero 1978) but, considering the millions of birds that fly between North and South America every year, some transport of seeds seems likely, especially in communities such as those of seacoasts frequented by migratory birds, in which selfcompatible and autogamous plants such as A. chamissonis are dominants (Peinado et al. 2009). From 301 N northward, A. chamissonis spreads across the Mediterranean climate region of Baja California and California. North of 421, in the temperate region, A. chamissonis populations always inhabit sites whose BIO is temperate submediterranean or Mediterranean (in Vancouver Island). In North America, Ambrosietea chamissonis is represented by three orders. Abronietalia maritimae and Abronietalia latifoliae group the vegetation dominated by perennial herbs forming shadow dunes when it colonizes the upper beach, and building discontinuous hillocks and hummocks when thriving on the seaward edge of dunes. On halfstabilized backdunes, the communities of both orders are progressively being replaced by fruticose communities of the new order Ambrosio chamissonis Eriogonetalia latifolii. Abronietalia maritimae groups the foredune associations from the Southern Californian and Martirense phytogeographical provinces (i.e. from Morro Bay southwards). These foredunes are broken up into separate conical hillocks (about 1.5 m in average height) created by the sand-stilling qualities of A. maritima and other perennial herbs with woody taproots. Unlike the northern communities of A. latifoliae, dune grasses are almost completely absent from this southern order. The main features of the order A. latifoliae and its unique alliance A. chamissonis, which group associations from Morro Bay northwards, arise from the sand-stilling abilities of the tap-rooted Abronia latifolia and two rhizomatous grasses, the native L. mollis, and the introduced Ammophila arenaria. Except for the communities of Ambrosion chamissonis, most of the Ambrosietea chamissonis associations in California correspond to the Sand-verbena Beach bursage series (Sawyer & Keeler-Wolf 1995), and to the Ambrosion chamissonis alliance of Pickart & Barbour (2007). Sawyer and Keeler-Wolf s series includes dune communities closest to ocean ones dominated by A. chamissonis and sand-verbenas (Abronia species), as well as inland communities dominated by chamaephytes and nanophanerophytes that we assign to the order Ambrosio chamissonis Eriogonetalia latifolii. The highly invasive behaviour of some introduced species of Aizoaceae, such as Carpobrotus chilensis, Carpobrotus edulis and Conicosia pungioniformis is causing the replacement of native Ambrosietea species in many places of California, mainly from Point Reyes southwards. The communities of A. chamissonis massively invaded by Aizoaceae correspond to Sawyer and Keeler- Wolf s Iceplant series. The communities of Ambrosietea chamissonis closely correspond to the Native Foredune Grassland (Pickart & Barbour 2007) and Sawyer and Keeler-Wolf s Native dunegrass series. As in temperate foredunes, much of this alliance has been reduced and replaced north of San Francisco by the naturalization of European grass A. arenaria, which is considered by Sawyer and Keeler-Wolf as an independent series: European beachgrass series. Native communities of Ambrosietea chamissonis created discontinuous often isolated, low hummocks, that rose gradually perpendicular to the shoreline aligned with prevailing NW onshore winds (Barbour & Johnson 1988). Although such native foredunes are still preserved in some places, most of them have been replaced by steep, continuous foredunes created by Ammophila (occurring at cover values of up to 99%), which may reach heights of several metres giving rise inland to a series of ridges and swales parallel to the shoreline. The dune scrub of Ambrosietea chamissonis Eriogonetalia latifolii, which closely corresponds to the Isocoma menziesii Lupinus chamissonis Ericameria ericoides shrubland alliance (Sawyer & Keeler-Wolf 1995), is endemic to Mediterranean California. It spreads from San Diego County northward to Bodega Bay. Except for some localities where Lupinus arboreus has been introduced, from Bodega northward evergreen shrubs such as Arctostaphylos columbiana, A. uva-ursi, Gaultheria shallon, Garrya elliptica or Vaccinium ovatum form sparse populations on backdunes under Pinion contortae forests. Although the Mediterranean rainfall pattern also exists north of Bodega Bay, precipitations are generally much greater than in the rest of California and summer drought is compensated by coastal fogs that supply an additional amount of unrecorded water of up to 200 mm, and by extended periods of cloudiness and fog which greatly reduce evaporation (Peinado et al. 2007). Both factors could be the cause of limiting the northward distribution of Ambrosio chamissonis Eriogonetalia latifolii. Two zones of vegetation encompass dune scrub: middunes and hind-dunes. The former are half-stabilized dunes transitional to foredune hillocks. Primary hillocks or foredune ridges built by pioneer species of Abronietalia maritimae or Abronietalia latifoliae are secondarily invaded by pioneer species of dune scrubs on foredune backs. Once stabilized by those plants, hind dunes support more complex communities thriving in deeper soils with a layer of litter or duff. In addition to this successional sequence visible in the larger dune systems, there are also clear variations among dune scrubs found on windward, more exposed slopes, when compared with their counterparts 10 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America on leeward, more stable slopes. The shrub element is poorly represented on windward slopes, where many pioneer species of Ambrosietea chamissonis and Cakiletea maritimae occur as scattered individuals colonizing the dune-scrub gaps. Dune scrub on lee slopes is denser, with a nearly closed canopy of shrubs. In addition to these floristic changes according to the autoecological requisites of the scrubs (see Barbour & Johnson 1988: Table 7.8, and Pickart & Barbour 2007: 164 to 168), the phytogeographical ranges of the main dune scrubs permit the distinction of two phytosociological complexes: a northern complex constituted by taxa ranging from northern California southward to San Diego County, and a southern complex, constituted by many Baja Californian endemics, which corresponds to the xerohalophytic class Atriplici julaceae Frankenietea palmeri, living on alkaline sandy soils (Peinado et al. 2008). The northern complex is well-characterized by a set of psammophilous shrubs and sub-shrubs endemic to the Southern Californian and Northern Californian provinces of the Californian region. We assigned those dune scrubs to the new order Ambrosio chamissonis Eriogonetalia latifolii, which encloses four alliances: Artemision pycnocephalae, Lupinion arboreo-chamissonis, Ericamerion ericoidis and Erigeronto glauci Eriophyllion staechadifolii. Habitats occupied by A. pycnocephalae, whose associations corresponds to the Artemision pycnocephalae alliance (McBride & Stone 1976; Pickart & Barbour 2007), are ecotones between the herbaceous vegetation of foredunes and the typical dune scrubs on backdunes. There is a gradual transition from dune mats dominated by Artemision pycnocephalae to dune scrub, with pioneering herbs (mainly A. chamissonis) able to colonize relatively unstable places on mid-dunes, and with pioneering shrubs (mainly Lupinus chamissonis and A. pycnocephala) able to colonize foredune backs. This is the ecologically intermediate position occupied by the communities of Artemision pycnocephalae in the psammosere, which is the same position occupied on the Mediterranean coasts by the suffrutescent communities of Crucianelletalia maritimae. In fact, the associations of Artemision pycnocephalae physiognomically resemble the European thermo- Atlantic grey dunes dominated by the suffrutescent C. maritimae on more or less stabilized soils low in humus. Lupinion arboreo-chamissonis includes dune scrubs on half-stabilized mid-dunes, that closely corresponds to several Lupinus chamissonis dune scrubs described by Williams & Potter (1972), Holton & Johnson (1979), and Barbour & Johnson (1988), and nitrophilous dune shrubs on disturbed backdunes, which correspond to the Lupinus arboreus Scrub (Heady et al. 1988), a widespread vegetation type whose greatest abundance coincides with plots having the highest values of per cent nitrogen and organic matter (Holton & Johnson 1979). Ericamerion ericoidis encloses denser dune scrubs on entirely stabilized hind dunes and lee slopes, which are comprised of a nearly closed canopy of E. ericoides under which abundant litter accumulates, that physiognomically resemble European heaths on decalcified dunes (Empetrion nigri, Calluno- Ulicetea Genistion pilosae and Ericion tetralicis), and correspond to the Haplopappus ericoides dune scrub described in many Californian dune systems (Barbour & Johnson 1988; Pickart & Barbour 2007). Erigeronto glauci Eriophyllion staechadifolii groups northern coastal scrubs on sea bluffs and on exposed slopes facing the ocean. Tropical beach and dune vegetation (Tables S10.3 and S10.4) The vegetation dominated by perennial plants, mainly by tap-rooted, succulent perennials, rhizomatous grasses and prostrate hemicryptophytes and chamaephytes, which act as pioneer colonizers of beaches and foredunes builders from tropical Baja California to Mesoamerica, belong to the class Euphorbio leucophyllae Sporoboletea virginici (Peinado et al. 2008). In the Tropical climate zone of Baja California, backdune communities are floristically well-defined. Dune scrubs of the Mediterranean association Loto bryanthi Isocometum menziesiii occur only in the northeastern tropical submediterranean zone, between southern El Vizcaíno and the dune complex of the Magdalenan barrier islands. In the tropical Sanlucan province Maytenus phyllanthoides dominates two backdune associations: Cyrtocarpo edulis Maytenetum phyllanthoidis, thriving on the leeward of the backdunes whose windward slopes are occupied by dense and flagged communities of Lycio congesti Maytenetum phyllanthoidis. The sandy inland plains immediately behind these dune systems are inhabited by columnar cacti and many small deciduous, sarcocaulescent trees. These sarcocaulescent thornscrubs growing on Arenosols form the association Jatropho cordatae Cyrtocarpetum edulis, which is floristically related to the psammophilous thornscrub climax of the association Antigono leptopi Cyrtocarpetum edulis (Peinado et al. 2008). Temperate and northern Mediterranean forested backdunes (Tables S10.5 and S10.6) The final succession stage on the coastal dunes of Oregon, Washington and in the forested dunes of Humboldt Bay and Tolowa in northern California comprises Picea sitchensis (on wet sands) or Pseudotsuga menziesii var. menziesii (on dry sands), which develop from different Pinus contorta var. contorta woods. According to their autoecological requirements, the five dune coniferous forest associations (Table 1: alliance 21) occupy different edatopes of dune zonation and represent different successional stages (see Peinado et al. 2011b). Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 11

Coastal vegetation of western North America Peinado, M. et al. On moist soils coniferous forests develop from different Salix thickets, which especially occur at the ecotone between moving dunes and forests (Wiedemann 1966, 1984; Kumler 1969; Franklin & Dyrness 1988; Pickart & Barbour 2007). The closed canopy of those dense thickets results from the dominance of two willows with two distinct ranges: Salix lasiolepis var. bigelovii, which thrives between Baja California and northern California, and Salix hookeriana, an essentially temperate species, which appears from British Columbia to the northwestern corner of California. Only there, north of Ten Mile, and in the adjacent southernmost part of the Oregon coast, do both willows intermix and many specimens show hybrid features. However, all willows on dune swales north of Ten Mile have been identified as S. hookeriana (references at Pickart & Barbour 2007: 160). The species making up the shrub layer of these thickets (1.5 4 m height) may become impenetrable and the ground under them devoid of plant life with only a layer of plant debris and leaves; an understory of sedges (Carex obnupta), rushes and other perennial herbs occurs in the usually more open thickets, but the herbaceous layer decreases as the canopy increases and in the tallest and thickest stands they flourish only on fringes and gaps. We included the Mediterranean willow thickets within the association Morello californicae Salicetum bigelovii, while those dominated by Hooker willows are ascribed to two associations: Lonicero ledebourii Salicetum hookerianae (Temperate Submediterranean) and Lonicero involucratae Salicetum hookerianae (Temperate Oceanic), both within the new class Salicetea lasiandro-exiguae. Early references to S. lasiolepis thickets in California are documented by Ramaley (1918) who described a willow thicket association with the dominance of this willow and Rubus ursinus, and an understory of two rushes: Juncus falcatus and Juncus leuseurii (a misapplied denomination for Juncus breweri in most early floras). There are swales with S. lasiolepis at Tomales Bay, Point Reyes, and Bodega Bay (Pickart & Barbour 2007) as well as at many other Californian localities (pers. obs.) but they are not regarded as distinct series by Sawyer & Keeler-Wolf (1995). Instead, Sawyer & Keeler-Wolf s Manual describes a Hooker willow series on the coastal dunes of northern California (1995: 262). Although S. lasiolepis thickets do not exist in Mediterranean Baja California, they are abundant as the understory of riparian sycamore forest dominated by Populus racemosa and in willow thickets along streams and rivers. Vegetation on temperate and Mediterranean deflation plains and dune swales (Table S10.7) Dune swales are seasonal or permanent wetlands that form in the troughs between dune ridges or in the deflation plain behind moving dunes. The bottoms of the wettest dune swales, which are always flooded, support freshwater marshes with Typha latifolia and Scirpus californicus where drainage off of the dunes provides sufficient fresh water. In areas transitional to salt marshes, where fresh and saline waters intermix, brackish marshes develop with species such as Carex lyngbyei, Juncus lesueurii, Schoenoplectus acutus and Schoenoplectus maritimus. Where saline waters predominate, pure salt marshes develop. These permanent, fresh, brackish or saline marshes are not dealt with here. Our analysis focuses on dune swales and on low level areas on deflation plains whose bottoms may be more or less waterlogged in winter and completely (or almost completely) dry during summer, and in which a zonation of herbaceous and woody communities develops because of a gentle gradation from the very wet, lowest bottoms, to the higher and drier margins. Differences of a few centimetres in ground surface elevation relative to the water table progressively separate distinct communities. Whereas in the isolated dune swales that appear encircled by dune mounds zonation is easily appreciable, on the deflation plains plant communities tend to form complex mosaics over the dune landscape. The herbaceous vegetation of these oligotrophic more or less wet habitats, usually dominated by rhizomatous species, mainly sedges and rushes, are here assigned to the new class Juncetea breweri. In this class, we include humid dune supporting communities ecologically related to the European class Isoëto Littorelletea, and grasslands and related short-lived, herb-rich plant communities of sandy, skeletal, and drought-stressed soils that thrive in the driest habitats of the deflation plains. In turn, these include communities ecologically and physiognomically related to the European class Koelerio Corynephoretea, and even share species such as: Aira caryophyllea, Aira praecox, Brachythecium albicans, Cerastium arvense, Ceratodon purpureus, Cladonia spp., Festuca rubra, Hypochaeris radicata, Racomitrium canescens, Rumex acetosella and Vulpia myuros. According to the moisture degree, two clearly recognizable new alliances of herbaceous communities occur: Argentino egedii Caricion obnuptae and Lupino littoralis Polygonion paronychiae. The wettest bottoms support very dense (average cover 4 80%) and almost pure communities of slough sedges (C. obnupta) reaching a height of 1 m, thriving on sites that are quite wet most of the year. On the temperate dunes, Wiedemann (1984) described a Carex obnupta Potentilla pacifica Community (P. pacifica = Argentina egedii), which is almost entirely restricted to very waterlogged habitats, where water persists 4 6 months of the year. In northern California, references to communities dominated by C. obnupta are limited to some plots and transects on herb 12 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America hollows analysed at Humboldt Bay (references in Pickart & Barbour 2007: 160). Moving away from the wettest bottoms, small changes in microtopography give rise to slightly more elevated and drier zones, where communities of small rushes (20 30 cm high) occur. These associations, dominated by two subspecies of Juncus falcatus (subsp. falcatus on Mediterranean swales, and subsp. sitchensis on temperate ones) accompanied by J. phaeocephalus, form meadows with scarce, scattered individuals of other species. The temperate association Juncetum breweri sitchensis correspond to the Rush Meadow Community, which occurs when water persists on the surface for 3 or 4 months during the winter (Wiedemann et al. 1969). References to communities related to the Mediterranean association Juncetum breweri falcati in northern California are restricted to four studies conducted on the Ten Mile dunes that were summarized by Pickart & Barbour (2007: 160). On soils that remain flooded only a few weeks after rains, communities dominated by the small sedge Carex pansa (height of its tallest specimens o 0.20 cm) develop. Despite its relative abundance, we are unaware of any reports of C. pansa communities, except for the Monterey Peninsula dunes, where McBride & Stone (1976: 123) described a Carex pansa community on dry swales. The new alliance Lupino littoralis Polygonion paronychiae closely corresponds to Wiedemann s Dry Meadow Community, which thrives in habitats where the water table is more than 91 cm below the surface during the summer months, and water never remains on the surface during the wet winter months (Wiedemann et al. 1969). These meadows were once dominated only by native species such as Festuca rubra, Glehnia leiocarpa, J. breweri, Lupinus littoralis, Poa macrantha, Polygonum paronychia or Solidago spathulata, which are today accompanied by A. arenaria in deteriorating stands. Conclusions The psammophilous coastal vegetation of the northern Pacific shows a distribution linked to the four MB (boreal, temperate, Mediterranean and tropical) that appear across the latitudinal gradient examined. This type of azonal vegetation follows patterns and shows limits of distribution that closely resemble those of the bioclimatically determined zonal vegetation. Generally, main physiognomic types are preferentially linked to a MB, although certain factors, mainly those related to soil moisture, determine that some psammophilous types, especially abundant in a particular MB, also appear in adjacent zones of different MB (bioclimatic ecotones). The four MB are characterized by floristic differences, which include the presence of endemic elements as well as elements of wider distribution, allowing us to establish relationships with the floras of Eurasia, South America and other continents. Each MB, in addition to its floristic peculiarities, shows both common and specific types of psammophilous vegetation spanning from associations to classes. Among the 12 phytosociological classes in which the study area s psammophilous vegetation is included, only two (C. maritimae and Stellarietea mediae), both linked to human disturbances, show a cosmopolitan distribution. A third, Honckenyo Elymetea arenarii, displays a Holarctic distribution and in the study area appears in the northernmost, coldest zones of boreal and temperate MB. Of the remaining nine classes, only one (Euphorbio leucophyllae Sporoboletea virginici) is exclusively linked to the tropical MB. The distribution of the communities of the further four classes is essentially linked to a given MB, but they also appear in ecotones of adjacent climate zones: (1) Ambrosietea chamissonis, which is exclusively Mediterranean but also exists in Temperate submediterranean areas; (2) psammophilous coniferous forests of the class Tsugetea mertensiano-heterophyllae, which are basically linked to the summer rains that characterize the temperate MB, but also penetrate the northern, wetter extreme of the Mediterranean zone, where fogs compensate summer droughts; (3) the drought-deciduous shrub and succulent communities of the class Prosopido torreyanae - Fouquierietea splendentis, which are widespread across all North American deserts of Tropical Arid bioclimate, but also appear in the small Mediterranean arid bioclimate zone of the El Vizcaíno desert in Baja California; and (4) the winter-annual communities of Achyronichio cooperi Abronietea villosae, which are more abundant in the Mediterranean arid bioclimate zone but also appear in some tropical areas of the Baja California Pacific, slightly influenced by winter rains. Another three classes mainly occur in a particular MB, but owing to their dependence on soil factors these also appear in climatically close areas: (1) Atriplici julaceae Frankenietea palmeri, whose distribution is mostly Mediterranean but its floristic ecological characterization is based on the presence of several Baja Californian endemisms exclusively linked to alkaline sands; (2) also linked to alkaline yet periodically flooded soils is the class Allenrolfeetea occidentalis, which is mainly linked to the study area s tropical arid bioclimate but may also sporadically appear in hyperarid Mediterranean zones of Baja California; and (3), the class Jucetea breweri, which is well represented in the temperate MB zone, but its mesophytic associations dominated by sedges and rushes also enjoy the wet sandy soils of the Mediterranean zone. Finally, the edaphically conditioned willow thickets living on dune swales (Morello californicae Salicetalia; Salicetea lasiandro-exiguae) are well-represented in the temperate and Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 13

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Ecological Studies 171, Springer, Berlin, DE. Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 15

Coastal vegetation of western North America Peinado, M. et al. Wiedemann, A.M., La Rea, D.J. & Smith, F.H. 1969. Plants of the Oregon coastal dunes. O.S.U. Book Store Inc., Corvallis, OR, US. Wiggins, I.L. 1980. Flora of Baja California. Stanford University Press, Stanford, CA, US. Wildi, O. 2010. Data analysis in vegetation ecology. Wiley- Blackwell, Sussex, UK. Williams, W.T. & Potter, J.R. 1972. The coastal strand community at Morro Bay State Park. Bulletin Torrey Botanical Club 99: 163 171. App. 1: Nomenclatural statement Arranged in alphabetical order, and following ICPN rules, this appendix includes nomenclatural types and diagnoses for syntaxa in need of typification. Diagnoses are completed by the descriptions made in the previous sections, especially in Table 1. Synonyms are followed by an indication of the ICPN articles, which lead to the delegation of a name into synonymy. Diagnoses of new associations and subassociations include: Table number and nomenclatural type relevé (holotypus): locality number as in Appendix S1; plot surface area (m 2 ); vegetation cover (%); list of species with the abundance/dominance value. For diagnostic species of associations see synoptic tables in Appendix S10 and phi-values in Appendix S11. Abronietalia latifoliae Knapp ex Peinado et al. ord. nov. hoc loco (Abronietalia latifoliae Knapp 1957 [art. 8]). Holotypus hoc loco: Ambrosion chamissonis in Plant Biosystems 135: 358 (2001). Diagnostic species: Abronia latifolia, Leymus mollis subsp. mollis (differential versus Abronietalia maritimae). Abronietum latifoliae Peinado et al. ass. nova hoc loco (Table S12.8). Holotypus: 155; 20; 60%. Abronia latifolia 4, Carpobrotus chilensis 1, Cakile maritima subsp. maritima1; ambrosietosum chamissonis Peinado et al. subass. nova hoc loco. Holotypus: 148; 20; 100%. Abronia latifolia 2, Ambrosia chamissonis 4, Carpobrotus chilensis 2, Fragaria chiloensis subsp. pacifica 3, Rumex salicifolius var. crassus 1;. leymetosum mollis stat. nov. hoc loco (Abronio latifoliae Elymetum mollis; holotypus: rel. 10 in Plant Biosystems 135: 356 [2001]). Achilleo arenicolae Lupinetum arborei Peinado et al. ass. nova hoc loco (Table S12.16). Holotypus: 178; 100; 80%. Lupinus arboreus 4, Achillea millefolium var. arenicola 2, Baccharis pilularis 1, Ammophila arenaria 1, Scrophularia californica 1, Solidago simplex subsp. simplex var. spathulata 1; eriophylletosum staechadifolii Peinado et al. subass. nova hoc loco. Holotypus: 167; 50; 100%. Lupinus arboreus 4, Achillea millefolium var. arenicola 2, Eriophyllum staechadifolium 3, Bromus carinatus var. maritimus 1. Agoserido eastwoodieae Lupinetum chamissonis Peinado et al. ass. nova hoc loco (Table S12.14). Holotypus: 167; 100; 80%. Lupinus chamissonis 4, Agoseris apargioides var. eastwoodieae 2, Bromus diandrus 1, Camissonia cheiranthifolia subsp. cheiranthifolia 1, Chorizanthe pungens 1, Erysimum menziesii subsp. concinnum1, Eschscholzia californica 1, Lupinus arboreus1, Marah fabaceus1, Poa douglasii 1, Polygonum paronychia 1, Pteridium aquilinum var. pubescens 2. Ambrosio chamissonis Abronietum maritimae Peinado et al. ass. nova hoc loco (Table S12.9). Holotypus: 127; 10; 80%. Abronia maritima 4, Ambrosia chamissonis 2, Camissonia cheiranthifolia subsp. cheiranthifolia 2, Calystegia soldanella 1; ambrosietosum chamissonis Peinado et al. subass. nova hoc loco. Holotypus: 136; 30; 80%. Abronia maritima 2, Ambrosia chamissonis 4, Carpobrotus chilensis 1; atriplicetosum leucophyllae stat. nov. hoc loco (Atriplici leucophyllae-abronietum maritimae; holotypus rel. 11 in Plant Biosystems 135: 356 [2001]). Ambrosio chamissonis Artemisietum pycnocephalae Peinado et al. ass. nova hoc loco (Table S12.10). Holotypus: 153; 20; 85%. Artemisia pycnocephala 5, Ambrosia chamissonis 3, Abronia latifolia 2, Carpobrotus chilensis1, Ammophila arenaria1, Leymus mollis subsp. mollis1. Ambrosio chamissonis Eriogonetalia latifolii Peinado et al. ord. nov. hoc loco. Holotypus: Artemision pycnocephalae alliancia nova. Diagnostic species: Achillea millefolium var. arenicola, Ambrosia chamissonis, Artemisia pycnocephala, Ericameria ericoides, Eriogonum latifolium, Ericameria ericoides, Eriophyllum staechaedifolium. Argentino egedii Caricetum obnuptae Peinado et al. ass. nova hoc loco (Table S12.28). Holotypus: 184; 20; 100%. Carex obnupta 4, Argentina egedii subsp. egedii 1, Carex pansa 2, Holcus mollis 1, Juncus breweri 1. Argentino egedii Caricion obnuptae Peinado et al. all. nova hoc loco. Holotypus: Argentino egedii-caricetum obnuptae Peinado et al. ass. nova. Diagnostic species. Argentina egedii subsp. egedii, Carex obnupta, Carex pansa, Juncus falcatus subsp. sitchensis, Veronica scutellata. Artemision pycnocephalae Peinado et al. all. nova hoc loco. Holotypus: Eriogono parvifolii-artemisietum pycnocephalae Peinado et al. ass. nova. Diagnostic species: Ambrosia chamissonis (differential versus Ericamerion ericoidis), Artemisia pycnocephala (dominant), Poa douglasii, Castilleja affinis subsp. affinis. Baccharido pilularis Ericamerietum ericoidis Peinado et al. ass. nova hoc loco (Table S12.17). Holotypus: 162; 50; 75%. Ericameria ericoides 4, Baccharis pilularis 2, Aira cariophyllea 1, Claytonia perfoliata subsp. mexicana1, Marah fabaceus1, Amsinckia spectabilis1. Cakiletum maritimo edentulae Peinado et al. ass. nova hoc loco (Table S12.7). Holotypus: 192; 50; 50%. Cakile edentula subsp. edentula var. edentula 2, Cakile maritima subsp. maritima 2, Leymus mollis subsp. mollis 1. Corethrogyno californicae Ericamerietum ericoidis Peinado et al. ass. nova hoc loco (Table S12.18). Holotypus: 132; 40; 80%. Ericameria ericoides 3, Corethrogyne filaginifolia var. californica 2, Croton californicus1, Conicosia 16 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America pugioniformis1, Eriogonum parvifolium 1, Lupinus chamissonis 2, Senecio blochmaniae 1. Cyrtocarpo edulis Maytenetum phyllanthoidis Peinado et al. ass. nova hoc loco (Table S12.24). Holotypus: 1; 200; 90%. Cyrtocarpa edulis 1, Maytenus phyllanthoides 4, Lycium fremontii var. congestum 2, Antigonon leptopus 2, Euphorbia leucophylla1, Jatropha cuneata 1, Pachycereus pringlei1, Jouvea pilosa 1. Dudleyo farinosae Ericamerietum ericoidis Peinado et al. ass. nova hoc loco (Table S12.19). Holotypus: 146; 50; 80%. Ericameria ericoides 4, Dudleya farinosa 1, Eriogonum latifolium 1, Polygonum paronychia 1. Ericamerion ericoidis Peinado et al. all. nova hoc loco. Holotypus: Dudleyo farinosae Ericamerietum ericoidis Peinado et al. ass. nova. Diagnostic species: Ericameria ericoides (dominant with cover 4 60%), Artemisia californica (differential versus Artemision pycnocephalae and Lupinion chamissonis). Erigeronto glauci Eriophylletum staechadifolii Peinado et al. ass. nova hoc loco (Table S12.21). Holotypus: 153; 50; 100%. Eriophyllum staechadifolium 5, Achillea millefolium var. arenicola 2, Erigeron glaucus 1, Eriogonum latifolium 1, Lupinus arboreus 1, Marah fabaceus 1, Sonchus oleraceus1. Erigeronto glauci Eriophyllion staechadifolii Peinado et al. all. nova hoc loco. Holotypus: Erigeronto glauci-eriophylletum staechadifolii Peinado et al. ass. nova. Diagnostic species: Angelica hendersonii, Eriophyllum staechadifolium (dominant with cover 4 60%), Erigeron glaucus, Rubus ursinus subsp. ursinus, Spergularia macrotheca, Stachys bullata. Eriogono parvifolii Artemisietum pycnocephalae Peinado et al. ass. nova hoc loco (Table S12.11). Holotypus: 146; 50; 90%. Artemisia pycnocephala 4, Corethrogyne filaginifolia var. californica1, Dudleya farinosa1, Ericameria ericoides1, Eriogonum latifolium 1, E. parvifolium 1, Eriophyllum staechadifolium 2, Lotus scoparius var. scoparius 1. Eriogono parvifolii Eriophylletum staechadifolii Peinado et al. ass. nova hoc loco (Table S12.20). Holotypus: 143; 50; 90%. Eriogonum parvifolium 2, Eriophyllum staechadifolium 3, Erigeron glaucus 1, Dudleya farinosa 1, Corethrogyne filaginifolia var. californica 1, Grindelia hirsutula var. maritima1, Artemisia pycnocephala 3, Carex pansa 1, Castilleja affinis subsp. affinis 1, Distichlis spicata 2. Ferocacto towsendiani Fouquierion diguetii Peinado et al. all. nova hoc loco. Holotypus: Burseretum hindsianomicrophyllae Peinado et al. in Journal of Vegetation Science 6(1): 88. 1995. Diagnostic species: Acacia goldmanii, A. peninsularis, A. virgata, Ambrosia magdalenae, Atamisquea emarginata, Bourreria sonorae, Bursera microphylla, Bursera odorata, Cardiospermum corindum, Condaliopsis rigida, Cyrtocarpa edulis, Euphorbia californica, Ferocactus towsendianus, Fouquieria diguetii, Jatropha cinerea, Jatropha cordata, Jatropha cuneata, Opuntia ciribe, Prosopis articulata, Ruellia californica, R. peninsularis, Tillandsia recurvata and Yucca valida. Heterotheco bolanderi Artemisietum pycnocephalae Peinado et al. ass. nova hoc loco (Table S12.12). Holotypus: 174; 40; 40%. Artemisia pycnocephala 2, Heterotheca sessiliflora subsp. bolanderi 2, Corethrogyne filaginifolia var. californica 1, Juncus breweri 1, Poa macrantha 1. Jatropho cordatae Cyrtocarpetum edulis Peinado et al. ass. nova hoc loco (Table S12.22). Holotypus: 17; 100; 70%. Cyrtocarpa edulis 3, Jatropha cordata 2, Jouvea pilosa 2, Proboscidea althaefolia 1, Bursera odorata 1, Oenothera drummondii var. thalassaphila 1, Prosopis palmeri 2, Dalea divaricata subsp. anthonyi 2. Juncetalia breweri Peinado et al. ord. nov. hoc loco. Holotypus: Lupino littoralis Polygonion paronychiae Peinado et al. all. nova hoc loco. Diagnostic species: See Juncetea breweri. Juncetea breweri Peinado et al. classis nova hoc loco. Holotypus: Juncetalia breweri ord. nov. Diagnostic species: Juncus breweri, Juncus falcatus subsp. sitchensis, Veronica scutellata. Juncetum breweri falcati Peinado et al. ass. nova hoc loco (Table S12.30). Holotypus: 162; 20; 80%. Juncus falcatus subsp. falcatus 4, Juncus breweri 1, Pseudognaphalium californicum 1, Polypogon monspeliensis 1, Argentina egedii subsp. egedii 1, Aira cariophyllea 1, Pseudognaphalium stramineum1, Carex obnupta1, Aira praecox1, Rumex salicifolius var. crassus 1. Juncetum breweri sitchensis Peinado et al. ass. nova hoc loco (Table S12.31). Holotypus: 195; 10; 60%. Juncus falcatus subsp. sitchensis 3, Juncus breweri 1, Hypochaeris radicata 1, Agrostis stolonifera1, Parentucellia viscosa 1, Pseudognaphalium purpureum1, Spiranthes romanzoffiana 1, Trifolium arvense 1, Trifolium campestre 1, Trifolium repens 1. Junco breweri Caricetum pansae Peinado et al. ass. nova hoc loco (Table S12.32). Holotypus: 186; 10; 80%. Carex pansa 3, Juncus breweri 2, Achillea millefolium var. arenicola1, Armeria maritima subsp. californica1, Aira cariophyllea 1, Festuca rubra subsp. rubra 1, Poa macrantha 1, Polygonum paronychia 1, Veronica scutellata1. Lathyro maritimi Leymetum mollis Peinado et al. ass. nova hoc loco (Table S12.4). Holotypus: 192; 50; 100%. Leymus mollis subsp. mollis, 5, Lathyrus japonicus var. maritimus, 2,Ammophila arenaria, 1,Erechtites minima, 1, Rubus ursinus subsp. ursinus1, Angelica hendersonii, 1, Agrostis stolonifera1, Rumex crispus1; Lathyretosum littoralis Peinado et al. subass. nova hoc loco. Holotypus: Locality: 147; 50; 70%. Lathyrus littoralis 3, Leymus mollis subsp. mollis 1, Cakile maritima subsp. maritima 2, Carpobrotus chilensis 2, Artemisia pycnocephala 1. Lathyro palustris Leymetum mollis Peinado et al. ass. nova hoc loco (Table S12.3). Holotypus: 259; 50; 100%. Lathyrus palustris 1, Leymus mollis subsp. mollis 5, Festuca rubra subsp. rubra 1, Hordeum brachyantherum subsp. brachyantherum 1, Poa macrocalyx 1, Calamagrostis canadensis1. Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 17

Coastal vegetation of western North America Peinado, M. et al. Leymo mollis Ammophiletum arenariae Peinado et al. ass. nova hoc loco (Table S12.6). Holotypus: 195; 100; 100%. Ammophila arenaria 5, Lathyrus japonicus var. maritimus 1, Leymus mollis subsp. mollis 1, Baccharis pilularis 1. Leymo mollis Caricetum macrocephalae Peinado et al. ass. nova hoc loco (Table S12.5). Holotypus: 238; 50; 80%. Carex macrocephala 3, Leymus mollis subsp. mollis 1, Ambrosia chamissonis1, Armeria maritima subsp. californica 1, Atriplex patula1, Cardionema ramosissimum 2, Rumex acetosella 1. Lonicero involucratae Salicetum hookerianae Peinado et al. ass. nova hoc loco (Table S12.27). Holotypus: 254; 100; 100%. Salix hookeriana 5, Carex obnupta 2, Lonicera involucrata var. involucrata 2, Morella californica 1, Scrophularia oregana 1, Plantago lanceolata1. Lonicero ledebouri Salicetum hookerianae Peinado et al. ass. nova hoc loco (Table S12.26). Holotypus: 185; 100; 100%. Salix hookeriana 5, Carex obnupta 2, Lonicera involucrata var. ledebourii 1, Rubus ursinus subsp. ursinus 1. Lupinion arboreo chamissonis Peinado et al. all. nova hoc loco. Holotypus: Senecioni blochmaniae Lupinetum chamissonis Peinado et al. ass. nova. Diagnostic species: Lupinus chamissonis, Lupinus arboreus, Phacelia ramosissima var. austrolitoralis. Lupino littoralis Polygonion paronychiae Peinado et al. all. nova hoc loco. Holotypus: Poo macranthae Lupinetum littoralis Peinado et al. ass. nova. Diagnostic species: Ammophila arenaria and Polygonum paronychia, (differential versus Argentino egedii-caricion obnuptae), Lupinus littoralis, Poa macrantha, Tanacetum camphoratum. Lycio congesti Maytenetum phyllanthoidis Peinado et al. ass. nova hoc loco (Table S12.23). Holotypus: 40; 200; 100%. Maytenus phyllantoides 5, Lycium fremonti var. congestum 2, Suaeda nigra 2, Abutilon carterae 1. Lycio Maytenion phyllanthoidis Peinado et al. all. nova hoc loco. Holotypus: Cyrtocarpo edulis Maytenetum phyllanthoidis Peinado et al. ass. nova. Diagnostic species: Lycium fremonti var. congestum, Maytenus phyllanthoides (cover 4 60%). Mertensio maritimae Honckenyetum majoris Peinado et al. ass. nova hoc loco (Table S12.1). Holotypus: 223; 20; 40%. Honckenya peploides subsp. major 2, Leymus mollis subsp. mollis1, Mertensia maritima var. maritima 1, Lathyrus japonicus var. maritimus 1, Atriplex gmelinii var. alaskensis 1. Morello californicae Salicetalia Peinado et al. ord. nov. hoc loco. Holotypus: Morello californicae Salicion alliancia nova. Diagnosis species: Lonicera involucrata var. involucrata, Morella californica, Salix hookeriana, Salix lasiolepis var. bigelovii, Scrophularia oregana. Morello californicae Salicion Peinado et al. all. nova hoc loco. Holotypus: Morello californicae-salicetum bigelovii Peinado et al. ass. nova. Diagnosis species: See Morello californicae Salicetalia. Morello californicae Salicetum bigelovii Peinado et al. ass. nova hoc loco (Table S12.25). Holotypus: 134; 100; 100%. Salix lasiolepis var. bigelovii 5, Morella californica 2, Toxicodendron diversilobum 1. Opuntio chollae Stenoceretalia gummosae Peinado et al. ord. nov. hoc loco. Holotypus: Idrio columnaris Pachycormion pubescentis Rivas-Martínez in Itinera Geobotanica 10: 98. 1997. Diagnostic species: Acacia brandegeana, Agave shawii subsp. goldmaniana, Ambrosia camphorata, Ambrosia carduacea, Ambrosia bryantii, Bursera hindsiana, Cochemia poselgeri, Dudleya ingens, Eriogonum trichopes, Euphorbia tomentulosa, Euphorbia xanti, Ferocactus fordii, Harfordia macroptera, Jatropha cuneata, Opuntia cholla, Opuntia echinocarpa var. nuda, Opuntia molesta, Opuntia prolifera, Opuntia rosarica, Opuntia tesajo, Pachycereus pringlei, Pedilanthus macrocarpus, Solanum hindsianum, Stenocereus gummosus, Vizcainoa geniculata and Yucca valida. Antigonum leptopus, Cyrtocarpa edulis, Lycium fremonti var. congestum and Maytenus phyllanthoides are differential species versus Parkinsonietalia florido microphyllae. Poetum douglasii Peinado et al. ass. nova hoc loco (Table S12.13). Holotypus: 167; 8; 70%. Poa douglasii 3, Agoseris apargioides var. eastwoodieae 1, Eschscholzia californica 1, Bromus carinatus 1, Pseudognaphalium purpureum1, Plantago subnuda1. Polygono paronychiae Tanacetetum camphorati Peinado et al. ass. nova hoc loco (Table S12.34). Holotypus: 185; 20; 50%. Tanacetum camphoratum 3, Polygonum paronychia 1, Cardionema ramosissimum 1, Armeria maritima subsp. californica 2; camissonietosum cheiranthifoliae Peinado et al. subass. nova hoc loco. Holotypus: 185; 20; 50%. Tanacetum camphoratum 2, Polygonum paronychia 2, Juncus breweri 2, Fragaria chiloensis subsp. pacifica 1, Camissonia cheiranthifolia subsp. cheiranthifolia 1, Poa macrantha 1. Poo macranthae Lupinetum littoralis Peinado et al. ass. nova hoc loco (Table S12.33). Holotypus: 199; 50; 40%. Lupinus littoralis 2, Poa macrantha1, Polygonum paronychia 1, Ammophila arenaria1, Hypochaeris radicata 1, Fragaria chiloensis subsp. pacifica1, Rumex acetosella 1, Festuca rubra subsp. rubra1, Pseudognaphalium purpureum1, Pinus contorta var. contorta1, Aira cariophyllea 1, Cytisus scoparius 1, Vaccinium ovatum1, Achillea millefolium var. littoralis1. Salicetalia delnortensis breweri Peinado et al. ord. nov. hoc loco. Holotypus: Salicion breweri Sánchez-Mata et Barbour in Barbour, Keeler-Wolf & Schoenherr (Terrestrial Vegetation of California, pp. 96, University of California Press, Berkeley, CA, US, 2007). Diagnostic species: Frangula tomentella, Salix breweri, Salix delnortensis. Salicetea lasiandro exiguae Peinado et al. classis nova hoc loco. Holotypus: Morello californicae Salicetalia Peinado et al. ord. nov. hoc loco. Diagnostic species: Salix bebbiana, Salix exigua, Salix laevigata, Salix lasiolepis var. lasiolepis, Salix lucida subsp. lucida,andsalix lucida subsp. lasiandra. 18 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America Senecioni blochmaniae Lupinetum chamissonis Peinado et al. ass. nova hoc loco (Table S12.15). Holotypus: 132; 100; 100%. Lupinus chamissonis 5, Achillea millefolium var. arenicola 2, Senecio blochmaniae 2, Corethrogyne filaginifolia var. californica 2, Phacelia ramosissima var. austrolitoralis1, Baccharis pilularis 1. Senecioni pseudoarnicae Leymion mollis Ohba, Miyawaki et Tx. ex Peinado et al. all. nova hoc loco (Carici macrocephalae-leymion mollis Ohba, Miyawaki & Tx. 1973 [article 3b]). Holotypus hoc loco: Senecioni pseudoarnicae- Leymetum mollis Peinado et al. ass. nova hoc loco. Diagnostic species: Achillea millefolium var. borealis, Conioselinum gmelinii, Honckenya peploides subsp. major, Lathyrus japonicus v. maritimus, Leymus mollis subsp. mollis, Ligusticum scoticum subsp. hultenii and Senecio pseudoarnica. Senecioni pseudoarnicae Leymetum mollis Peinado et al. ass. nova hoc loco (Table S12.2. Holotypus: 260; 100; 100%. Leymus mollis subsp. mollis 4, Senecio pseudoarnica 2, Conioselinum gmelinii 2, Angelica lucida1, Ligusticum scoticum subsp. hultenii 2, Honckenya peploides subsp. major1, Galium aparine 1, Epilobium ciliatum subsp. glandulosum 1, Heracleum maximum 2. Veronico scutellatae Caricetum obnuptae Peinado et al. ass. nova hoc loco. (Table S12.29). Holotypus: 192; 20; 100%. Carex obnupta 5, Argentina egedii subsp. egedii 2, Veronica scutellata 1, Juncus falcatus subsp. sitchensis 1, Lotus corniculatus1, Salix hookeriana1. Supporting Information Additional Supporting Information may be found in the online version of this article: Appendix S1. Figure S1: Sampled localities. Appendix S2. Figure S2: Physiographic map of the North American Pacific basin and phytogeographical sketch of the study area. Appendix S3. Strand and dune forms, main features, major vegetation types, geological and geomorphological processes. Appendix S4. Sampled localities, bioclimates, potential natural vegetation, ecofloristic zones, zonobiomes, zonoecotones, ecoregions and phytogeographical provinces. Appendix S5. Climate data and indices, bioclimatic classification, zonobiomes and potential natural vegetation at the association level for 226 weather stations. Appendix S6. Distribution ranges of the phytogeographical elements and recorded taxa. Appendix S7. Dendrograms obtained from nine ALC. Appendix S8. Main features of the eight data sets prepared for the second ALC. Appendix S9. Relevé groups obtained through nine successive ALC. Appendix S10. Synoptic syntaxonomical tables. Appendix S11. Diagnostic species and phi-values for higher syntaxa and associations. Appendix S12. Complete phytosociological tables for new associations. Appendix S13. Dichotomous key to associations based on floristic, phytogeographical and ecological features. Plate S1. Beach and dune plants: 1 = Abronia latifolia. 2=Abronia umbellata subsp. umbellata. 3=Mimulus guttatus. 4=Glehnia littoralis subsp. leiocarpa. 5=Astragalus fastidius. 6=Euphorbia leucophylla. 7=Rhododendron albiflorum. 8=Ambrosia chamissonis. 9=Carex macrocephala (male, left; female, right). 10 = Chaenactis lacera. 11 = Scaevola plumieri. 12 = Stegnosperma halimifolium. 13 = Lonicera involucrata var. ledebourii (inflorescence). 14 = Lonicera involucrata var. ledebourii (berry). 15 = Camissonia crassifolia. 16 = Houstonia mucronata. 17=Lathyrus japonicus var. maritimus. 18=Senecio pseudoarnica. 19=Commelina diffusa. 20=Lotus bryantii. 21 = Picea sitchensis. 22 = Pseutotsuga menziesii var. menziesii. 23 = Pinus muricata. 24 = Pinus contorta var. contorta. 25 = Portulaca pilosa. 26 = Atriplex leucophylla. 27 = Antigonon leptopus. 28=Polygonum paronychia. 29=Coulterella capitata. 30 = Drymaria holosteoides var. crassifolia. Photographs 1 4, 7 9, 13, 14, 21 24, 26 and 28 by L. Monje, Scientific Drawing & Photography Department, Alcalá University (LMO). Plate S2. (a) Lonicero ledebourii Pinetum sitchensis and freshwater swamp with Nuphar luteum subsp. polysepalum on Croft Lake, Oregon. Photograph by LMO. (b) Arctostaphylo uva-ursi Pinetum contortae on dune ridges. Bare sands colonized by Lupino littoralis Polygonion paronychiae with Arctostaphylos uva-ursi (1) and Cytisus scoparius (2). Arctostaphylos columbiana (3) at forest edges. Umpqua River, Oregon. (c) Wind-trained Picea sitchensis and krummholzs with Gaultheria shallon, Garrya elliptica and Lonicera ledebourii. Sutton Beach, Oregon. Photo by LMO. (d) Isolated remnant of dune parabolas still bearing forest (Lonicero ledebouri Pinetum contortae) resting upon the summit at Cape Kiwanda, Oregon. Photo by LMO. (e) Poo macranthae Lupinetum littoralis, with Poa macrantha and Juncus breweri in the foreground, on dry deflations plains among young shore pines (Pinus contorta). Background: Arctostaphylo uva-ursi Pinetum contortae. Nehalem Bay State Park, Oregon. (f) Leymo mollis Caricetum macrocephalae among driftwoods. Sand Lake, Oregon. Photo by LMO. (g) Detail of Juncetum breweri sitchensis. Dominants are Juncus falcatus subsp. sitchensis (flat leaves) and Juncus breweri (with lateral inflorescences and leafless). Bullard Beach, Oregon. (h) Foreground: (1), Mertensio maritimae Honckenyetum majoris on debris line. Background: (2) Senecioni pseudoarnicae-leymetum mollis. Middle Bay, Kodiak Island, Alaska. Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 19

Coastal vegetation of western North America Peinado, M. et al. Plate S3. Baja California. (a) Coastal flat, dominated by Atriplici linearis Frankenietum palmeri, near Laguna San Ignacio, Baja California Sur. (b) Euphorbio miserae Lycietum californici on windy alkali sandy plains near Bahía Asunción, Baja California Sur. (c) Foredune built by Suaedo taxifoliae Allenrolfeetum occidentalis near Mulegé, Baja California Sur. Backdunes with Lycio congesti Maytenetum phyllanthoidis (2). (d) Pebble beach with Atriplicetum magdalenae bordering debris lines near Punta Abreojos, Baja California Sur. (e) Dense communities of Lycio congesti Maytenetum phyllanthoidis thriving on windward slopes of the Sanlucan backdunes. (1) Maytenus phyllanthoides; (2) Lycium congestum. Punta Conejo, Baja California Sur. (f) Sarcocaulescent thornscrubs: (1) Cyrtocarpo edulis Maytenetum phyllanthoidis on leeward slopes of the Sanlucan backdunes; (2) Jatropho cordatae Cyrtocarpetum edulis on inland sandy plains. Between Punta Colorada and Punta Arena, Baja California Sur. (g) Foredune hillocks of Palafoxio linearis Abronietum maritimae in Isla Magdalena, Baja California Sur. Visually, dominants are Abronia maritima (green) and Sesuvium verrucosum (red). Dominant NW winds coming from the right side of the picture. (h) Shadow dunes and small hillocks of Euphorbio leucophyllae Jouveetum pilosae on the upper part of the beach, and pioneer vegetation (Euphorbio leucophyllae Drymarietum crassifoliae) along the tidal line. Near Punta Arenas, Baja California Sur. (i) Detail of Plate 4, photo E. (1),Active dunes of the Flandrian Episode II with Senecioni blochmaniae Lupinetum chamissonis; (2) older, stabilized, parabola of the Flandrian Episode I at bottom, with Corethrogyno californicae Ericamerietum ericoidis; (3) dominant on the foreground are Lupinus chamissonis (blue flowers), Eriophyllum staechadifolium (yellow inflorescences) and Achillea arenicola (white inflorescences). (j) Interior of the tip of a Flandrian parabola at Marina State Beach, California. Foreground: wind-exposed flank with Polygono paronychiae Artemisietum pycnocephalae (1) and wind-sheltered depression with Dudleyo farinosae Ericamerietum ericoidis (2). Background: Preflandrian dune sheets with irrigated crops and remnants of the climax. Photograph by LMO. (k) Wind-exposed top of the former parabola. Dune mat (Polygono paronychiae Artemisietum pycnocephalae). Dominants are (1) Artemisia pycnocephala (canescent), (2) Polygonum paronychia and Eriogonum parvifolium (prostrate chamaephytes) and Dudleya caespitosa (yellow flowers). Photograph by LMO. l = Lee slope of the former parabola with Dudleyo farinosae Ericamerietum ericoidis. Foreground: (1) Eriogonum latifolium (white, rounded inflorescences) and Dudleya farinosa; (2) Ericameria ericoides; Background: (3) windexposed crest with Polygono paronychiae Artemisietum pycnocephalae); (4) top of a dune invaded by Carpobrotus species. Photograph by LMO. Plate S4. (a) Dune swale at Umpqua River dune system, Oregon. (1) Pioneer clumps of Ammophila arenaria on active sands; (2) Marshy bottom with helophytes such as Eleocharis palustris and Juncus phaeocephalus, encircled by sedge and rush meadows (Argentino egedii-caricion obnuptae); (3) sedge meadows (Argentino egedii Caricetum obnuptae) with scattered Salix hookeriana seedlings; (4) Willow thickets (Lonicero Salicetum hookerianae); (5) Canopies of semi-buried willows emerging from (6) hummocky areas created Leymo mollis Ammophiletum arenariae and Lupino littoralis Polygonion paronychiae; (7) Carici obnuptae Pinetum contortae on wet sands; (8) Isolated remnant of dune parabolas bearing forest (Lonicero ledebouri Pinetum contortae) resting upon the summit. Photograph by LMO. (b) Flat-topped dune and rear inland plains between Punta Colorada and Punta Arena, Baja California Sur. (1) Euphorbio leucophyllae Drymarietum crassifoliae forming small mounds along tidal lines; (2) Euphorbio leucophyllae Jouveetum pilosae building a first ridge on the upper part of the beach; (3) second level of the ridge (Ipomoeo imperati Jouveetum pilosae); (4) third level of the ridge (Daleo anthonynae Jouveetum pilosae); (5) thornscrub (Cyrtocarpo edulis Maytenetum phyllanthoidis) thriving on leeward slopes; (6) sandy plains with sarcocaulescent thornscrubs (Jatropho cordatae Cyrtocarpetum edulis). (c) Dry deflation plain at Nehalem Bay State Park, Oregon. Foreground and open areas: Lupino littoralis Polygonion paronychiae with scattered Arctostaphylos uva-ursi and Pinus contorta var. contorta seedlings. Background: Arctostaphylo Pinetum contortae. (d) Flandrian dunes perched on a steep bluff rising 100 m from the beach (BE) at Mussel Rock. Dominant NW winds coming from the left corner of the picture. Active dunes of the Flandrian Episode II (E-II) invading an older, stabilized and eroded, parabola of the Flandrian Episode I (E-I). E-II: Ambrosio chamissonis Abronietum maritimae ambrosietosum chamissonis. E-I: Corethrogyno californicae Ericamerietum ericoidis. Beach: Atriplex leucophylla forming shadow dunes in the upper beach zone and Cakile maritima on debris lines. Photograph by LMO. (e) Mediterranean dune complex near Guadalupe, California in a picture taken with the ocean around 2.5 km behind. Foreground: active dunes of the Flandrian Episode II (E-II) invading an older, stabilized, parabola of the Flandrian Episode I (E-I). E-II: Senecioni blochmaniae Lupinetum chamissonis. E-I: Corethrogyno californicae Ericamerietum ericoidis, secondarily invaded by Lupinus chamissonis and by the exotic, very aggressive, grass Ehrharta calycina. Background: stabilized Preflandrian dune sheets (PF), active Flandrian dunes of Mussel Rock (MR; see picture D), and Casmalia Hills. Photo by LMO. Photo S5. Pholisma arenarium (Lemnoaceae), an extremely rare root parasite, endemic to the North 20 Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science

Peinado, M. et al. Coastal vegetation of western North America American Southwest, always living on coastal dunes and sandy soils. In this photograph, taken near Morro Bay, California, occurs as parasitic on Ericameria ericoides, a diagnostic species of the association Corethrogyno californicae-ericamerietum ericoidis. Photo by Luis Monje, Scientific Drawing & Photography Department, Alcalá University. Please note: Wiley-Blackwell is not responsible for the content or functionality of any supporting materials supplied by the authors. Any queries (other than missing material) should be directed to the corresponding author for the article. Doi: 10.1111/j.1654-109X.2011.01134.x r 2011 International Association for Vegetation Science 21