Phytogeography of Hevenor Inlet and surrounds on Pitt Island, BC Adam Huggins

Supervisor: Dr. Geraldine Allen University of Victoria Biology 490A 11/30/2017

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20 to Bella Coola in the south. Thus, the vast Introduction majority of the terrain can only be accessed by boat, aircraft, or foot. The North Coast of BC Pitt Island The North Coast of (BC) is home to some of the most rugged and Pitt Island, at 1,373 km2 , is the fifth largest inaccessible terrain in . island in BC (Scott, 2009), and is underlain by Extending roughly 500 km from the northern syn- and post- terrane accretion plutonic tip of Vancouver Island to the -BC rocks of the Captain’s Cove suite - primarily border in western Canada, this region is tonalite (Crawford et al., 1999). Named by characterized by the highest mean annual Captain George Vancouver after William Pitt, precipitation in North America and consists of the British PM at the time of his voyage, the a series of mountain ranges, flooded valleys, island falls within the traditional and unceded and coastal islands (Klock & Mullock, 2001). A territory of the Git lax m’oon or Gitxaała First defining feature of this landscape is the Nation, a Sm'álgyax-speaking people within relatively recent withdrawal of the the Tsimshian group (Menzies, 2016). Cordilleran ice sheet (CIS) at the onset of the Currently, there is no human habitation on Holocene, resulting in an impoverished flora the island, although small fishing vessels composed primarily of species that have frequent certain areas and small hunting recently colonized the region from elsewhere cabins are present. Logging and mining (Ogilvie & Ceska, 1984; Peinado et al., 2009). operations are ongoing in the region, but are Bounded by the Coast Ranges on the east and not immediately underway in the vicinity of by the Hecate Strait and Haida Gwaii on the the study area, located up Hevenor Inlet. west, most of the region is blanketed in dense evergreen forests and bogs and falls within Hevenor Inlet (Figure 1) is a long oceanic inlet the Coastal Western Hemlock (CWH) or about two-thirds of the way up the island that Mountain Hemlock (MH) biogeoclimatic provides convenient marine access to its zones, the former below 600m and the latter rugged alpine interior. The presence of old above 600m (Radcliffe et al., 1994). It is village sites, fish traps, canoe pullouts, and sparsely populated and served by only three estuarine root gardens along the shoreline of main roads: Highways 16 and 37 to Prince the inlet confirm former habitation and use of Rupert and Kitimat in the north, and Highway this area by Gitxaała; in addition, unpublished

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ethnographic accounts indicate that alpine Channel (Nagorsen & Keddie, 2000); or resources, including mountain goats persistence following rapid post-glacial (Or eamnos americanus) and useful , insular colonization from southerly, northerly, and/or cryptic refugial populations, for which there exists fossil (ibid.) and genetic evidence (Shafer et al, 2010a). The lack of data for the Pitt Island population precludes any conclusions at this point. A third, unexplored hypotheses - that the population was transplanted from mainland populations by Tsimshian hunters long ago - was one of several primary motivating factors for the field work that gave rise to this paper. Figure 1: Overlooking Hevenor Inlet on Pitt Island, BC were harvested annually by Gitxaała people Postglacial Colonization here in the early 20th century (C. R. Menzies, Pitt Island was completely blanketed by the personal communication, June 28 2017). The CIS during the Last Glacial Maximum (LGM), study area lies at the head of Hevenor Inlet which occurred between 25,000 and 20,000 and extends northwards to Hevenor Peak, the years before present (BP) on the coast highest point on the island at 1099 m. (Lacourse et al., 2012). Deglaciation in the

Hecate lowland, where Pitt Island is located, Perhaps the most notable feature of the proceeded in fits and bursts starting, island is its small population of mountain minimally, at 12,700 BP and terminating by goats, which present the only natural, extant 10,000 BP (Clague, 1984); however, the insular population on the northwest coast Hecate Strait and significant portions of Haida (Nagorsen & Keddie, 2000), although there Gwaii and the Alexander Archipelago of are now several introduced populations on Alaska were free of glacial ice between Alaskan Islands, including Baranof Island (but 15,000 and 18,000 BP (Ager et al., 2009; see Shafer et al., 2010a). There are presently Lacourse et al., 2012). It is likely that portions two hypotheses to explain this anomalous of Pitt Island were deglaciated somewhere population: infrequent immigration and between these two time ranges, although supplementation from Kitimat Range isostatic fluctuations in sea level populations across the 0.5 km Grenville

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Figure 2: Geographic location of study area in coastal BC relative to putative cryptic refugia and CIS extent at LGM. Based on Shafer et al., 2010b. may have inundated some of these areas prior refugia in Beringia (Brubaker et al., 2005; to 8,000 BP, when present-day sea levels were DeChaine, 2008) and the Pacific Northwest established (Clague, 1984). (Shafer et al., 2010b). In recent years, phylogeographic and paleoecological support At the onset of the Wisconsin glaciation, for the presence of cryptic refugia on the species retreated north or south to ice-free western margin of the CIS has grown (ibid.;

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Dixon, 2013). The Alexander Archipelago (Carrera et al., 2007; Ager et al., 2009), Haida Methods Gwaii and the Hecate Strait (Mathewes et al., 2015), and northern Vancouver Island (Ogilvie Field Work & Ceska, 1984) are all considered to have Field work on Pitt Island took place over four potentially supported refugial populations of consecutive expeditions between May and plants, animals, and insects (Shafer et al., July of 2017. Activities included cutting trails, 2010b). Pitt Island’s geographic position surveying for mountain goats, getting wet (Figure 2) places it centrally in relation to and drying off again, resetting wildlife these cryptic refugia and nearly equidistant cameras, and looking for signs of pre-contact between Beringian and Pacific Northwest human use. inventories and collections refugia. While it is extremely unlikely that it were peripheral tasks performed during spare supported glacial refugia during the LGM (but time. In all, 51 specimens were see Roberts & Hamann et al., 2015), Pitt collected, and 137 vascular plants were Island’s proximity to cryptic refugia and identified and noted. Field identifications remoteness from northerly and southerly were made with reference to Pojar and refugia makes it an excellent case study of MacKinnon (2016). Specimens were pressed in insular postglacial recolonization on the BC lightweight, homemade presses, and relevant coast. data, including flowering dates at varying

altitudes, was recorded in a field notebook. In This paper explores the dynamics of the addition, 196 observations of around 119 postglacial recolonization of the North Coast species were uploaded into an iNaturalist of BC by examining the phylogeography of project, which is viewable at the vegetation of Pitt Island. It combines https://www.inaturalist.org/projects/hevenor data from 51 new collections, an informal -inlet-study. The GPS coordinates of these inventory of 137 species, and an analysis of observations are scrambled on the public herbarium collections from this region. A pages to discourage would-be hunters from discussion of the development of regional retracing our steps. floral biogeographic elements, using examples from Pitt Island, will follow. Collections

All specimens were keyed with reference to Hitchcock and Cronquist (1973), Cody (1996),

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and Hultén (1968); modern binomials were Prince Rupert Forest Region) and the Great assigned with according to Klinkenberg Bear Rainforest, but omit the Portland Inlet (2017). Collections will be housed at the and include northern parts of the Central University of Victoria Herbarium. Coast. Haida Gwaii and Vancouver Island are excluded due to their unique geographic Inventories contexts and glacial histories. Collections that were georeferenced within this area but The Pitt Island vascular plant field inventory were clearly collected elsewhere were was conducted informally on “trails” crossing excluded. very nearly the full contingent of site series present on the island, from the shoreline of Hevenor Inlet to the peak of Hevenor Mountain. It is by no means complete, especially with regards to graminoid species. In addition, a thorough inventory of vascular plants was performed on Hevenor Islet, a tiny rock protrusion with rich soils and an impressive collection of species located near the mouth of Hevenor Inlet. Figure 3: Boundaries of the search zone and locations of

collections from the Consortium of Pacific Northwest After field work was complete, a regional Herbaria database. inventory was prepared. A search was performed for all vascular plant collections The data was transferred to a spreadsheet, from the north coast of BC on the Consortium duplicate species entries were eliminated, of Pacific Northwest Herbaria database, and obsolete taxons were updated. Next, yielding 2,524 specimens. The boundaries of EFlora BC, the USDA plants database, the search zone (Figure 3) were roughly as CalFlora, and the Flora of North America were follows: Greaves Island and Owikeno Lake to cross-referenced to determine habitat and the south; the Alaska-BC border to the north; distribution for each species. The the Hecate strait to the west; and Kitimat and distributions were each categorized within a Bella Coola to the east. These boundaries generalized North American floristic element: loosely conform to the boundaries of the these include Cosmopolitan (C), North North Coast Forest District (a district of the American (NA), Western (W), West Coast (CW), Pacific Northwest (PNW), coastal Pacific

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Northwest (CPNW), northern North American communities recorded in a field notebook. As (NNA), northern and western North American thorough SMR and SNR analyses were not (NNAW), and coastal British Columbian (CBC) performed, these assignments are provisional distributions. Additionally, species with and are provided to inform future studies. amphiberingian, circumboreal, or disjunct ranges were noted. See Appendix D for a Results visual representation of these distributions. The number of species within each of these Collections floristic elements was tallied and then calculated as a percentage of the total. The These collections, to my knowledge, are the same calculation was performed for the Pitt first in the study area and the first on Pitt Island inventory for comparison purposes. Island since those made in June 1979 by W. B. Schofield. In all, 51 specimens were identified Site Series Classification to (sub)species - these are listed in Appendix A. Many of the specimens represent species Pitt Island’s vegetation communities fall that are very common regionally, but endemic within the Coastal Western Hemlock Very Wet to the northwest coast: C arex circinata, Hypermaritime (CWHvh2) BGC zone below Ranunculus cooleyae, L igiusticum calderi, around 600 m elevation, the Mountain Lupinus nootkatensis, C optis aspleniifolia, Hemlock Wet Hypermaritime windward Pedicularis ornithorhyncha, G entiana (MHwh1) BGC zone between 600 and 900 m douglasiana, and F ritillaria camschatcensis a re elevation, and the as-yet undescribed examples. Others are amphiberingian - G eum Mountain Hemlock Wet Hypermaritime calthifolium, Ha rrimanella stelleriana, a nd parkland (MHwh1p) and Alpine Tundra (AT) Carex macrochaeta, for example - and still BGC zones above about 900 m elevation others are more broadly distributed in North (Radcliffe et al., 1994). Slope and America and elsewhere: examples include groundwater table are the main determinants Triantha glutinosa, E mpetrum nigrum, and of vegetation type regionally, with Eriophorum angustifolium. unforested areas tending towards paludification (Asada et al., 2003). Individual A few specimens represent range extensions. site series were assigned to the areas visited Populations of P enstemon davidsonii, with reference to Banner et al. (1993) based discovered on rock outcrops along a lone on field observations of vegetation ridgeline in the study area, are well north of

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previous collections, including those on Haida amphiberingian distribution, 7.46% have Gwaii. Robust populations of A nemone significant disjunct populations (primarily on narcissiflora a nd G entiana platypetala a re the southern tip of South America), and only among the southernmost collected, with 2.02% are cosmopolitan. exception to northern Vancouver Island, which may have been a refugial area (Ogilvie With regards to distribution within North & Ceska, 1984). L uzula arcuata unalaschcensis, America, 58.87% of the species are confined in particular, h ad not been collected in coastal to the western half of North America, with BC outside of northern Vancouver Island. almost half of these – 25.60% of the total – occurring only in the Pacific Northwest; many North Coast Inventory of these are also amphiberingian. Slightly less than half, or 41.13%, of the species occur Overall, 567 species are represented by the across northern North America; many of these 2,524 specimens from the Consortium of are circumboreal. Just 4.84% are found only Pacific Northwest Herbaria. Of these, 71, or along the coast of the northern Pacific 12.42%, were recently introduced from Northwest, and a majority of these are also Eurasia, Africa, Australia, South America, or amphiberingian, with several being obligate eastern North America. The remaining 496 coastline species of circumboreal distribution. species are considered native and are used for subsequent calculations. Species numbers Pitt Island Inventory and percentages for various floristic elements can be found in Table 1; broader distribution 137 species were noted in the informal attributes are considered in Table 2. The inventory of the study area. In a noticeable inventory is included in Appendix B. departure from the trends captured by the North Coast inventory, not a single one of Several trends are worth noting. First, almost these species is known to have been exactly half (48.4%) of the species are found introduced. Species numbers and only in North America, while the other half percentages for various floristic elements can (51.6%) are found in North America and be found in Table 1; broader distribution elsewhere in the world. The most significant attributes are considered in Table 2. The trend is that over a half, or 27.42% of the inventory is included in Appendix C. total, of these species have a circumboreal Trends in this inventory mirror the regional distribution. Of the total, 14.72% have an trends from the North Coast inventory, with a

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Table 1: Comparison of Pitt Island inventory and North Coast inventory species and percent of total falling within identified floristic elements. Percentage of introduced species taken using the total; other percentages are taken using the total minus introduced species. States / Provinces are abbreviated, with parentheses indicating placement at the periphery of the floristic element.

Floristic Description Pitt Island Pitt Island North Co. North Co. Element Species Percent Species Percent C Cosmopolitan 4 2.92% 10 2.02% NA Throughout North America 15 10.95% 80 16.13% W North America, w of the Rocky Mtns. 27 19.71% 107 21.57% CW (Baja CA), CA, w OR and w W, BC, (AK) 16 11.68% 43 8.67% (N CA), (OR), WA, BC, AK, (ID), (AB), PNW (MO), (NV) 14 10.22% 60 12.10% CPNW (N CA), (w OR), w WA, w BC, AK 28 20.44% 58 11.69% NNA North America, n of ~40° 23 16.79% 69 13.91% North America, n of ~40° and incl. areas NNAW w of the Rocky Mtns. 2 1.46% 45 9.07% CBC (w WA), W BC (endemic), (AK) 8 5.84% 24 4.84% I Introduced 0 0.00% 71 12.52% TOTAL 137 567

Table 2: Comparison of Pitt Island inventory and North Coast inventory species and percent of total with additional notable distribution attributes. Percentages are taken using the total from above minus introduced species.

Distribution Description Pitt Island Pitt Island North Co. North Co. Attribute Species Percent Species Percent a Amphiberingian 23 16.79% 73 14.72% cb Circumboreal / Circumpolar 35 25.55% 136 27.42% ic Confined to the immediate coast 8 5.84% 27 5.44% d Disjuncts in other ranges 9 6.57% 37 7.46% slightly higher (67.88%) proportion of species being confined to western North America. Site Series Classification This is likely due to the elevated percentage Site series for the study area are shown in of species confined to the coastal Pacific Figure 4. These assignments are provisional Northwest and lower percentages of species and require more thorough on-site sampling. found throughout most of North America in High elevation assignments are limited by this versus the regional inventory. their placement in undescribed BGC units.

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Figure 4: Map of the Hevenor Inlet study area, showing paths traversed during fieldwork, provisional site series designations, landscape features, notable plant collections, mountain goat sightings, and indirect observations of mountain goat territory. Unnumbered site series are from undescribed BGC zones. BGC units in standard format.

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There are two significant departures between Discussion the inventories. First, there is a slightly higher proportion of species confined to the coastal Marr et al. (2012) describe northern BC as a Pacific Northwest (CPNW) in the Pitt Island contact zone between five floristic elements: inventory than in the regional inventory, with circumpolar, amphiberingian, Yukon-Alaska- a reciprocal higher proportion of broadly northwest B.C. endemics, North American distributed North American (NA, NNAW) radiants, and Cordilleran groups. The local species in the regional inventory than in the and regional inventories, while admittedly Pitt Island inventory. This may be due to the coarse, are mutually reinforcing and capture close proximity of Pitt Island to putative several of these critical phylogeographic cryptic refugia, discussed later. Second, trends within the regional flora. These can be roughly 12% of the total in the regional summarized as follows: inventory are introduced species, while not a ❖ Roughly half of the species present in single introduced species was noted in the the region are endemic to North study area on Pitt Island. This can be America; the other half are shared attributed the infrequency of anthropogenic with other continents. This trend is disturbance on Pitt Island. corroborated by Peinado et al. (2009).

❖ Around 60-70% of the species occur The relative lack of regional floral endemism, only west of the Rocky Mountains in noted elsewhere (Peinado et al., 2009), is a North America; of these, about half direct result of recurrent glaciation in the are restricted to the Pacific recent geologic history of the North Coast of Northwest. BC. The flora of Pitt Island, and the North ❖ A quarter of the species are Coast in general, is composed of species that circumboreal. weathered these “ice ages” in select refugia: ❖ 15% of the species are amphiberingian these include North America south of the ice ❖ Minor contingents of species are sheets, Beringia, and putative cryptic refugia confined to the immediate coastlines, on nunataks within the ice sheets and along or have disjuncts on other continents, the western margin of the CIS (Shafer et al., primarily southern South America. 2010b). These three refugial “zones” will be ❖ A small proportion of the species discussed below, using representative (<5%) is endemic to the North Coast. collections from Pitt Island for illustration.

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Montana and Colorado (ibid.). A number of From the North: Beringia these arctic-alpine lineages appear to have originated in Asia and dispersed from there A significant species contingent weathered (Marr et al., 2008; Marr et al., 2013; Allen et al. the recent glaciations in the dry, unglaciated 2012; Allen et al. 2015; Guest & Allen, 2014), expanse of northwestern Alaska and although considerable population structuring northeastern Siberia known as Beringia. within refugia can complicate analyses During the LGM, eustatic lowering sea levels (Brubaker et al., 2005; Allen et al., 2012). This exposed large portions of the continental reciprocal dispersal pattern is also shelf in this region, creating a land bridge responsible for the distribution of between Asia and North America (Brubaker et amphiberingian species such as Ha rrimanella al., 2005) that is widely considered to have stelleriana and C arex macrochaeta, the latter served as a bridge for species exchange of which appears to be retreating northwards between the continents (Dixon, 2013; as the climate warms (Wilson et al., 2008). DeChaine, 2008). Ancestors of indigenous

North Americans are thought to have crossed Beringia also served as a refugium for boreal this land bridge from Asia to North America forest species, including P inus contorta ssp. (Dixon, 2013). contorta, Picea spp. Populus spp., Betula spp.,

and L arix spp. (Brubaker et al., 2005; Roberts & Many of the species that recolonized BC from Hamann, 2015; Godbout et al., 2008). Of Beringia are characteristic of a circumboreal these, only P . contorta ssp. contorta i s found “arctic-alpine” flora (Marr et al., 2012) that is on Pitt Island. Other species that likely restricted to arctic regions during glacial dispersed south to Pitt Island from Beringian periods and expands southwards through refugia include V accinium uliginosum ( Eidesen alpine habitat corridors during warmer et al., 2007) and E mpetrum nigrum ( Popp et interludes. When ice sheets form following al., 2011). The latter species is one of several warm intervals, southerly populations of that have disjunct populations in the southern these species may be isolated but persist in hemisphere; this can be explained by chance high-altitude environments: A nemone avian long-distance dispersal events from narcissiflora i s an example of a species that is north to south (ibid.). largely confined to circumboreal latitudes

(and is found only infrequently on the coast Not all of the arctic-alpine species on Pitt south of Alaska) but has relict populations in Island came from Beringian refugia: in fact,

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many of these species appear to have Other refugial areas in North America include recolonized BC from populations both north the Rocky Mountains (Guest & Allen, 2014), and south of the CIS (Marr et al. 2008; Marr et and the Beartooth Plateau (Marr et al., 2013; Allen et al. 2012; Godbout et al., al., 2012), the Columbia Basin (Godbout et al., 2008; Guest & Allen, 2014; Shafer et al. 201a). 2008), and (Roberts & Hamann, Sibbaldia procumbens survived in North 2015). Species from these refugia primarily America south of the CIS and followed the ice colonized the interior of BC or became relicts. sheets north as they melted (Allen et al., Species and lineages in coastal BC mostly 2015), which is surprising given its current originated in refugia west of the Cascades range. The list of species considered to have and north of the Klamath Mountains (Shafer inhabited Beringian refugia (Shafer et al., et al., 2010b). Many of the species observed 2010b; Roberts & Hamann, 2015) is actually and collected on Pitt Island recolonized the notable for how few of its species were North Coast from southerly refugia: examples observed on Pitt Island. It is likely that the include P enstemon davidsonii, D odecatheon majority of species found on Pitt Island and jeffreyi, and C astilleja spp. along the North Coast of BC dispersed northwards along the coast from the south. From the West?: Cryptic refugia

Although fossil evidence is still lacking, From the South: North America genetic and circumstantial evidence has North America, south of the CIS, is best accumulated to suggest that refugia occurred considered not as a single refugium, but along the western margin of the CIS during rather a number of distinct refugia separated the LGM (Shafer et al., 2010b) and previous by north-south trending mountain chains glaciations (Mathewes et al., 2015). These (Shafer et al., 2010b; Roberts & Hamann, “cryptic refugia” are thought to have occurred 2015). For example, most of the tree species in the Alexander Archipelago of Alaska (Ager found on Pitt Island - including A bies amabilis, et al., 2010; Carrara et al., 2007), Haida Gwaii Picea sitchensis, Tsuga heterophylla, Tsuga and the Hecate Strait (LaCourse et al., 2012; mertensiana, Tsuga heterophylla, and Mathewes et al., 2015), and northern Xanthocyparis nootkatensis - display minimal Vancouver Island (Ogilvie & Ceska, 1984). genetic variation suggestive of a relatively Large sections of the continental shelf were small, isolated Pacific Northwest refugium exposed during the LGM in these areas, which during the LGM (Roberts & Hamann, 2015). were also the first to be deglaciated as the

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climate warmed. Favorable exposures on the western faces of these islands may have been ice-free even during the LGM (Ager et al., 2010; Carrara et al., 2007; Dixon, 2013). Some species, including mountain goats (Nagorsen & Keddie, 2000) and P inus contorta ssp. contorta (Godbout et al., 2008) were present and even widespread so soon after deglaciation (and were more common then than now) that dispersal from northerly or southerly refugia would have had to have been extremely rapid to make sense.

Pitt Island is unlikely to have served as a refugium during the LGM (but see Figure 5). Nevertheless, it is positioned centrally between the three best-supported putative cryptic refugia, and would have been rapidly colonized by any refugial species as the glaciers receded. There is genetic and circumstantial evidence to support refugial populations of P inus contorta ssp. contorta i n this region (Godbout et al., 2008; Roberts & Figure 5: Reconstructed glacial refugia at LGM based on Hamann, 2015), although these genetic GCM paleoclimate simulations and genetic data. Note reconstructions are contested by prior study the inclusion of Pitt Island within the Haida Gwaii - (Macdonald & Cwynar, 1985). Haida Gwaii, in Hecate Strait - Alexander Archipelago refugial zone. particular, is recognized for its unusual From Roberts & Hamann, 2015. endemic plant (and moss) species. L igusticum calderi i s a good example of a species that is Enemion savilei i s found only on Haida Gwaii, known to have been endemic to Haida Gwaii Brooks Peninsula on Vancouver Island, and but has since radiated eastward to the North Porcher Island (Pojar & MacKinnon, 2016) - Coast and Pitt Island (Mathewes et al., 2015). which is just north of Pitt Island. This unusual distribution is characteristic of a number of

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species - including L uzula arcuata and nonexistent outside of the coastal Pacific Gentiana platypetala, which were collected on Northwest - were observed to be dominant Pitt Island - which have been explicitly on Pitt Island, especially at higher elevations: referenced as evidence for glacial refugia these include L upinus nootkatensis, Gentiana (Ogilvie & Ceska, 1984). And, of course, the douglasiana, Gentiana platypetala, Ranunculus singular, anomalous presence and persistence cooleyae, Carex circinata, and Geum of mountain goats on Pitt Island cannot be calthifolium. These are the same species that discounted, especially given fossil (Nagorsen dominate similar areas on northern & Keddie, 2000) and genetic (Shafer et al., Vancouver Island (Ogilvie & Ceska, 1984). 2010a) evidence supporting cryptic refugia While evidence is lacking, the nearly for this species. monocultural dominance of these taxa on Pitt Island with respect to their apparently absent It is worth noting that a small handful of relatives and surrogates is suggestive of rapid species - many of which are uncommon or colonization by available species from nearby refugia. These observations, however, could also be partly explained by the dominance of extreme moisture regimes in determining the composition of regional plant communities (Asada et al., 2003).

Finally, there is some evidence to suggest that cryptic refugia might have existed on nunataks w ithin t he CIS in Northern BC during the LGM (Marr et al., 2008; Shafer et al., 2010a; Godbout et al., 2008). It would be extremely challenging to find fossil evidence to confirm this. Furthermore, these refugia probably have little bearing on the postglacial colonization of the North Coast or Pitt Island,

which are proximate to more established Figure 6: Postulated postglacial colonization routes, cryptic refugia. with blue arrows indicating primary pathways and yellow arrows indicating secondary pathways. From Shafer et al., 2010b.

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Human Intervention?

There is a growing body of evidence demonstrating that coastal First Nations communities were heavily involved in the active management of coastal ecosystems (Turner et al., 2013). Documented activities include(d) transplantation of species - Figure 7: F ritillaria camschatcensis h arvest on Pitt Island. especially F ritillaria camschatcensis ( Figure 7), which is locally abundant on Pitt Island in meadow habitats - to new locations, pruning, coppicing, tilling, and fertilization. While documentation of Gitxaała ecosystem management has been primarily focused on marine resources (Menzies, 2016), regionally important species such as F ritillaria camschatcensis, Malus fusca, T axus brevifolia, and other fruiting species were likely Figure 8: Hevenor Islet (center) in Hevenor Inlet beneath transplanted within the study area, where the mountainous study area, viewed facing due East. they tend to occur in accessible, highly visible locations. Hevenor Islet (Figure 8), in particular, has remarkably well-developed organic soils for a rocky protrusion of its size, and is vegetated almost exclusively by useful species (Figure 9), recalling the description of “orchard gardens” in Turner et al. (2013). This little outcrop is a prime fishing site and is a central feature of Hevenor Inlet, which was once a Gitxaała thoroughfare. A complete Figure 9: Human amongst M alus fusca, Rubus parviflorus, vascular inventory of Hevenor Islet is included Vaccinium alaskaense, Vaccinium ovalifolium, Vaccinium in Appendix E. Further study should elucidate parvifolium, Ribes laxiflorum, Sambucus racemosa, a nd the extent of regional cultural management. Polypodium glycyrrhiza o n Hevenor Islet.

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thought to be t he c orridor that facilitated Limitations and Future Directions human colonization of the Americas (Dixon, 2013), it is essential that archeological and Study of the postglacial colonization of BC, botanical exploration continue in the region. long stymied by a poor fossil record, has picked up in recent years with the adoption of Pitt Island presents many avenues for genetic analyses. While these methods have ongoing study. As a large, topographically revolutionized the fields of botany, biology, complex island located near proposed cryptic and paleoecology, they are not without refugia, it could provide a great deal of limitations. In particular, it has been noted valuable information about the history of that different sets of genetic markers within glaciation and deglaciation in the region. As a the same species can suggest different documented site of First Nations inhabitation conclusions; as such, it is best practice to and management, it could shed further light combine datasets (Eidesen et al., 2007). This on the histories and lives of the coast’s first dynamic can be glimpsed in the discrepancies inhabitants. Several lines of inquiry present between restriction fragment (Marr et al., themselves. First, simple botanical 2008) and cpDNA sequence (Allen et al., 2012) explorations like this would be beneficial to analyses of Ox yria digyna p hylogeography. In our understanding of the regional flora and addition, recurrent range expansions and likely turn up a few surprises. Second, genetic contractions, vicariance, and introgression can sampling of the local mountain goats (which complicate these analyses (Eidesen et al., are often ignored in larger studies) and 2007; Marr et al., 2008; Allen et al., 2015). comparison to data from throughout BC (see

Shafer et al., 2010a) might elucidate the A further limitation particular to the North history of this unusual population. Peat cores Coast of BC is its inherent inaccessibility and from Pitt Island, like those from Mitkof and the resulting paucity of collections in the Hippa Islands (Ager et al., 2009; Lacourse et region. A number of the species that were al., 2012), would clarify the postglacial collected on Pitt Island were not even chronology for this part of the ‘inner outer represented in the Consortium of Pacific coast.’ Genetic and morphological Northwest Herbaria search for the North examination of species at the periphery of Coast. Regional alpine environs, in particular, their range on Pitt Island, such as P enstemon are understudied, especially with respect to davidsonii and L uzula arcuata, could reveal human use. Given that the coast of BC is novel trends in regional phylogeography,

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much like those discussed in Marr et al. The flora of the North Coast of BC is relatively (2012). Finally, ongoing anthropological and depauperate due to a history of recurrent archaeological exploration of the island is glaciation (Peinado et al., 2009; Ogilvie & almost sure to turn up additional evidence of Ceska, 1984) and restrictive climatic human use through the millenia. conditions (Asada et al. 2003). It was deglaciated early on after the LGM and was Conclusion rapidly colonized, primarily from southerly refugia in present-day and The North Coast of BC is, in many respects, Oregon (Roberts & Hamann, 2015), but also one of North America’s final frontiers for from Beringian refugia in the north (Brubaker exploration. In recent years, it has captured et al., 2005) and cryptic refugia along the the attention and imagination of scientists margin of the CIS (Carrara et al., 2007; and nonscientists alike for its proposed role in Mathewes et al., 2015). Patterns of facilitating human colonization of the vegetation turnover appear consistent, if Americas (Dixon, 2013), as well as the unique temporally staggered, across the region (Ager species and subspecies that may have et al., 2009; Lacourse et al., 2012). Patterns of inhabited its refugia during glacial episodes vegetation on Pitt Island reflect these (Shafer et al., 2010b). Yet, it is also the home regional trends and provide circumstantial and backyard of coastal First Nations who are evidence in support of the presence of cryptic increasingly recognized as having developed refugia nearby during the LGM (Ogilvie & some of the most sophisticated and Ceska, 1984). Coastal First Nations likely successful systems of terrestrial and marine manipulated plant communities at specific resource management in the world (Turner et sites through transplantation and other al., 2013; Menzies, 2016). The informal management activities; it is also possible that inventories, collections, and observations these interventions had broader effects on discussed here would be of marginal value the regional flora (Turner et al., 2013). Future were they gathered nearly anywhere else in investigations should enrich our North America; however, given the relative understanding of regional phytogeography, paucity of field work on the North Coast, they cryptic refugia, colonization corridors, represent a valuable, if minor, addition to our traditional management activities, and the understanding of the history and antiquity of humanity in the Americas. phylogeography of the region.

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Acknowledgements

I am grateful to Dr. Charles Menzies and UBC Anthropology for making this field work possible. Charles was extremely generous with his knowledge and gave me the opportunity to explore and make collections in his ancestral territory even when there were many other important tasks to be completed. Thanks are also due to Ada Smith and Marion Travis, who rounded out our team. It was Marion’s idea to collect plants, and she spent many hours with me identifying species, making collections, and drying specimens. Special thanks to Dr. Geraldine Allen for supervising this project and assisting with specimen identification. Dr. Joe Antos provided assistance with Poaceae, and Rande Kane provided moral support. Finally, thanks to Heidi Guest and Rachel for preparing and mounting the specimens.

References

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Appendix A: Pitt Island Collections List

Table 3: List of numbered collections from Pitt Island included with this paper, with iNaturalist references.

Subspecies / Collection # Genus Species Variety iNat Reference 1 Moneses uniflora 6571785 2 Cochlearia groenlandica 6571795 3 Trientalis europaea arctica 4 Harrimanella stelleriana 6571921 5 Coptis aspleniifolia 6571804 6 Huperzia haleakalae 6571848 7 Empetrum nigrum 8 Ranunculus cooleyae 6572010 9 Gentiana douglasiana 6571926 10 Geum calthifolium 6572011 11 Trichophorum cespitosum 12 Listera caurina 6576676 13 Eriophorum angustifolium 14 Dodecatheon jeffreyi 6572014 15 Streptopus lanceolata 6950948 16 Pinguicula vulgaris 17 Pedicularis ornithorhyncha 6950941 18 Platanthera aquilonis 19 Carex macrochaeta 20 Eriophorum angustifolium 21 Anemone narcissiflora 6951562 22 Luzula arcuata unalaschcensis 6973956 23 Trientalis europaea arctica 6973963 24 Castilleja parviflora 6973919 25 Viola langsdorffii 6951659 26 Viola glabella 6951482 27 Tiarella trifoliata unifoliata 28 Castilleja miniata miniata 6951203 29 Fritillaria camschatcensis 6951171

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30 Stelleria crispa 6951133 31 Cardamine oligosperma kamtschatica 6951082 32 Lupinus nootkatensis nootkatensis 33 Micranthes nelsoniana carlottae 6951227 34 Claytonia sibirica 6951071 35 Carex mertensii 6956380 36 Carex macrochaeta 6956177 37 Leptarrhena pyrolifolia 38 Carex circinata 8747782 39 Arnica latifolia 6956185 canadensis / 40 Calamagrostis canadensis scabra 7319680

41 Erigeron peregrinus peregrinus 7319677 42 Ligusticum calderi 7319545 43 Triantha glutinosa 7319553 44 tolmiei 7321953 45 Penstemon davidsonii menziesii 7321891 46 Gentiana platypetala 7322009 47 Deschampsia elongata (possible hybrid) 7322062 48 Gentiana douglasiana 7321862 49 Epilobium anagallidifolium 6951058 50 Artemisia norvegica saxatilis 7319532 51 Saxifraga ferruginea 7319705

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Appendix B: Specimen inventory of the North Coast of BC

Table 4: Annotated vascular flora species list of the North Coast of BC from Consortium of Pacific Northwest Herbaria, with notes on floristic element, distribution attributes, and habitat. Floristic element and attribute abbreviations are described in Tables 1 and 2. Quotes (“) following a species element indicate that the range refers to the subspecies and was not considered in calculations.

Family Scientific Name Element Notes on range and habitat preferences

Widespread throughout N America; Adoxaceae Sambucus racemosa NAcb circumboreal

Sambucus racemosa var. arborescens (racemosa) Widespread throughout the coastal West Adoxaceae (pubens) CWcb" from CA north to AK; circumboreal

Sambucus racemosa var. Mostly SE BC and throughout E NA; Adoxaceae leucocarpa NA" probably planted

Primarily an interior species of the West; Sambucus racemosa var. rare and local on the BC coast, probably Adoxaceae melanocarpa W" planted

Frequent across northern N America from Adoxaceae Viburnum edule NNA the PNW to the NE

Atriplex dioica (Atriplex Shorelines and saline soils throughout N Amaranthaceae subspicata) NA America

Shorelines throughout the coastal PNW Amaranthaceae Atriplex gmelinii CPNW from OR to AK

Shorelines and saline soils throughout N Amaranthaceae Atriplex patula NAcb America; circumboreal

Beaches and marshes of the coastal West Salicornia pacifica from MX north to AK; also Atlantic and Amaranthaceae (Salicornia virginica) NAic Gulf coasts

Wet areas of the PNW from N CA north to Apiaceae Angelica arguta PNW S BC; probably planted

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Wet areas of the coastal PNW from N CA Apiaceae Angelica genuflexa CPNWa north to AK; amphiberingian

Wet areas of the coastal PNW from N CA Apiaceae Angelica lucida CPNWa north to AK; amphiberingian

Wet areas of the West from CA north to Apiaceae Cicuta douglasii W AK

Conioselinum gmelinii Wet areas of the lowland coastal West Apiaceae (Conioselinum pacificum) W from CA north to AK

Glehnia littoralis ssp. Coastal dunes and beaches of the coastal Apiaceae leiocarpa CWica West from CA north to AK; amphiberingian

Endemic to islands of coastal BC from Apiaceae Ligusticum calderi CBC Vancouver Island north to SE AK

Beaches and bluffs of coastal BC north to Apiaceae Ligusticum scoticum CBCa AK; amphiberingian

Ligusticum scoticum ssp. Beaches and bluffs of coastal BC north to Apiaceae hultenii CBCa" AK; amphiberingian

Wet areas of the West from CA north to Apiaceae Oenanthe sarmentosa W AK

Osmorhiza berteroi Widespread throughout the West from CA Apiaceae (Osmorhiza chilensis) Wd north to AK; disjunct in S America

Widespread throughout the PNW from N Apiaceae Osmorhiza purpurea PNW CA north to AK

Wet areas of N America; apparently Apiaceae Sium suave NA uncommon on north coast of BC

Wet areas of the PNW from OR north to Araliaceae Oplopanax horridus PNWd AK; disjunct in Great Lakes region

Frequent in the PNW from N CA north to BC; apparently uncommon on the north Aristolochiaceae Asarum caudatum PNW coast of BC

Widespread in the coastal PNW from N Asparagaceae Maianthemum dilatatum PNW CA north to AK

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Maianthemum Widespread throughout the West from CA racemosum ssp. north to AK; apparently uncommon on the Asparagaceae amplexicaule W north coast of BC

Widespread throughout N America; apparently uncommon on the north coast Asparagaceae Maianthemum stellatum NA of BC

Aspleniaceae Asplenium trichomanes NAcb Rocky areas of N America; circumboreal

Aspleniaceae Asplenium viride NAcb Rocky areas of N America; circumboreal

Widespread throughout N America and the Asteraceae Achillea millefolium NAcb Northern Hemisphere

Achillea millefolium ssp. North coast of BC north to AK; Asteraceae borealis CBCa" amphiberingian

Achillea millefolium var. Sporadic along the pacific coast of N Asteraceae pacifica CW" America

Beaches of the coastal West from CA Asteraceae Ambrosia chamissonis CWic north to AK

Asteraceae Anaphalis margaritacea NA Widespread throughout N America

Asteraceae Antennaria howellii NA Montane zones throughout N America

Widespread throughout the West from CA north to AK and across northern N Asteraceae Antennaria rosea NNAW America

Widely introduced throughout N America Asteraceae Arctium minus I from Eurasia

Wet areas of the West from CA north to Asteraceae Arnica amplexicaulis W AK

Arnica amplexicaulis ssp. Widespread in montane zones of the West Asteraceae amplexicaulis W" from CA north to AK

Arnica chamissonis ssp. Widespread in montane and alpine zones Asteraceae foliosa W of the West from CA north to AK

Widespread in montane and alpine zones Asteraceae Arnica cordifolia W of the West from CA north to AK

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Artemisia norvegica ssp. saxatilis (Artemisia Montane and alpine zones of the West Asteraceae arctica) Wcb from CA north to AK; circumboreal

Widespread throughout the West from CA Canadanthus modestus north to AK and across northern N Asteraceae (Aster modestus) NNAW America

Introduced throughout N America from Asteraceae Cirsium vulgare I Eurasia

Widespread throughout N America; Asteraceae Conyza canadensis I probably introduced

Erigeron acris var. Wet montane and subalpine zones across Asteraceae asteroides NNAcb northern N America; circumboreal

Interior PNW from OR north to SE BC; Asteraceae Erigeron corymbosus PNW probably planted

Widespread throughout montane and subalpine zones of the West from CA Asteraceae Erigeron peregrinus W north to AK

Widespread throughout montane and subalpine zones of the West from CA Erigeron peregrinus ssp. north to AK; mostly west of Asteraceae callianthemus W" Coast­Cascade mountains

Widespread throughout montane and Erigeron peregrinus ssp. subalpine zones of the West from CA Asteraceae peregrinus W" north to AK; common throughout BC

Introduced throughout N America from Asteraceae Hieracium aurantiacum I Eurasia

Montane and alpine zones of the West Asteraceae Hieracium gracile W from CA north to AK

Introduced throughout N America from Asteraceae Hieracium maculatum I Eurasia

Hieracium scouleri var. Widespread throughout the West from CA Asteraceae griseum W north to AK

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Subalpine and alpine zones of the West Asteraceae Hieracium triste Wa from CA north to AK; amphiberingian

Widespread across northern N America; Asteraceae Hieracium umbellatum NNAcb circumboreal

Introduced throughout N America from Asteraceae Hypochaeris radicata I Eurasia

Lactuca biennis (Lactuca Asteraceae spicata) NA Throughout N America

Leucanthemum vulgare (Chrysanthemum Introduced throughout N America from Asteraceae leucanthemum) I Eurasia

Introduced throughout N America from Asteraceae Matricaria matricarioides I Eurasia

Wet meadows and bogs of the coastal Asteraceae Microseris borealis CW West from CA north to AK

Nabalus alata Asteraceae (Prenanthes alata) CPNW Coastal PNW from OR north to AK

Widespread throughout the PNW from OR north to AK and across northern N Asteraceae Packera paupercula NNA America; also SE N America

Rare in interior BC; disjunct from eastern Asteraceae Packera plattensis NA N America; possibly planted?

Packera subnuda Montane and subalpine zones of the PNW Asteraceae (Senecio moresbiensis) PNW from N CA north to AK

Petasites frigidus ssp. Widespread throughout the West from CA Asteraceae nivalis Wcb north to AK; circumboreal

Beaches and dunes of extreme NW BC north to AK; amphiberingian; disjunct on Asteraceae Senecio pseudoarnica CBCadic the North Atlantic coast

Wet areas of the West from CA north to Asteraceae Senecio triangularis W AK

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Introduced throughout N America from Asteraceae Senecio viscosus I Eurasia

Introduced throughout N America from Asteraceae Senecio vulgaris I Eurasia

Widespread throughout the West from CA Solidago lepida var. north to AK and across northern N Asteraceae lepida NNAW America

Introduced throughout N America from Asteraceae Sonchus arvensis I Eurasia

Sonchus arvensis var. Introduced throughout N America from Asteraceae glabrescens I Eurasia

Symphyotrichum foliaceum (Aster Widespread throughout the West from CA Asteraceae foliaceus) W north to AK

Symphyotrichum spathulatum (Aster Widespread throughout the West from CA Asteraceae occidentalis) W north to AK

Symphyotrichum subspicatum (Aster Widespread throughout wet areas of the Asteraceae subspicatus) W West from CA north to AK

Introduced throughout N America from Asteraceae Tanacetum vulgare I Eurasia

Introduced throughout N America from Asteraceae Taraxacum officinale I Eurasia

Subalpine and alpine zones of N America; Athyriaceae Athyrium distentifolium NAcb circumpolar

Widespread in wet areas of N America; Athyriaceae Athyrium filix­femina NAa amphiberingian

Athyrium filix­femina var. Widespread in wet areas of N America; Athyriaceae cyclosorum NAa amphiberingian

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Coastal West from CA north to AK; Balsaminaceae Impatiens noli­tangere CWcb circumboreal

Wet areas of the (mostly) coastal West Betulaceae Alnus rubra CW from CA north to AK

Wet areas of the West from CA north to AK and across northern N America; Betulaceae Alnus viridis NNAWcb circumboreal

Alnus viridis ssp. crispa Betulaceae (Alnus crispa) CBC" Wet areas of N BC

Wet areas of the West from CA north to Alnus viridis ssp. sinuata AK and across northern N America; Betulaceae (Alnus sinuata) NNAWcb" circumboreal

Betulaceae Betula glandulosa NA Common throughout northern N America

Widespread in the coastal West from CA north to AK; circumboreal with disjuncts in Blechnaceae Blechnum spicant CWcbd N Africa and Eurasia

Beaches of the coastal West from CA Boraginaceae Amsinckia spectabilis CWic north to BC

Boraginaceae Hackelia deflexa NNAcb Northern N America; circumboreal

Interior PNW from OR north to SE BC; Boraginaceae Mertensia longiflora PNW faulty record?

Introduced throughout N America from Boraginaceae Myosotis discolor I Eurasia

Wet areas of the West from CA north to Boraginaceae Myosotis laxa Wcbd AK; circumboreal; also S America

Introduced throughout N America from Boraginaceae Myosotis scorpioides I Eurasia

Introduced throughout N America from Boraginaceae Myosotis sylvatica I Eurasia

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Arabidopsis lyrata ssp. kamchatica (Arabis Montane and subalpine zones of the PNW Brassicaceae kamchatica) PNWa from WA north to AK; amphiberingian

Arabis eschscholtziana Widespread throughout the PNW from OR Brassicaceae (Arabis hirsuta) PNW north to AK

Brassicaceae Barbarea orthoceras NA Widespread throughout N America

Introduced throughout N America from Brassicaceae Barbarea verna I Eurasia

Introduced throughout N America from Brassicaceae Barbarea vulgaris I Eurasia

Beaches and shorelines of the coastal West from CA north to AK; also shorelines Brassicaceae Cakile edentula CWd of the Great Lakes and Atlantic coast

Cakile edentula var. Beaches and shorelines of the coastal Brassicaceae californica CW" West from CA north to AK

Introduced to beaches of the coastal West Brassicaceae Cakile maritima I from Eurasia

Coastal PNW from N CA north to BC; known only from Vancouver Island and Brassicaceae Cardamine angulata CPNW Haida Gwaii in BC

Widespread throughout the West from CA Brassicaceae Cardamine oligosperma Wa north to AK; amphiberingian

Cardamine oligosperma Brassicaceae var. kamtschatica CBCa" Coastal BC north to AK; amphiberingian

Cardamine oligosperma Widespread throughout the West from CA Brassicaceae var. oligosperma W" north to S BC

Brassicaceae Cardamine pensylvanica NA Widespread throughout N America

Beaches, tidal marshes, and mudflats of Cochlearia groenlandica the coastal PNW from WA north to AK; Brassicaceae (Cochlearia officinalis) CPNWcbic circumboreal

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Draba hyperborea (Draba Shorelines of coastal BC north to AK; Brassicaceae grandis) CBCa amphiberingian

Lepidium virginicum var. Widespread throughout N America; Brassicaceae pubescens NA probably introduced

Introduced to wet areas throughout N Brassicaceae Nasturtium officinale I America from Eurasia

Rorippa curvipes var. Widespread throughout the West from CA Brassicaceae integra W north to BC

Widespread in wet areas throughout N Brassicaceae Rorippa palustris NAcb America; circumboreal

Rorippa palustris ssp. Widespread in wet areas throughout N Brassicaceae hispida NAcb" America; circumboreal

Rorippa palustris ssp. Widespread in wet areas throughout N Brassicaceae palustris (occidentalis) NA" America

Subularia aquatica ssp. Infrequent in wet areas throughout N Brassicaceae americana NA America

Widespread throughout N America; Brassicaceae Turritis glabra NAcb circumboreal; status uncertain

Widespread throughout N America; Campanulaceae Campanula rotundifolia NAcb circumboreal

Wet areas across northern N America; Campanulaceae Lobelia dortmanna NNAcb circumboreal

Caprifoliaceae Lonicera involucrata NA Widespread throughout N America

Caprifoliaceae Symphoricarpos albus NA Widespread throughout N America

Widespread in wet areas of the montane and subalpine zones of the West from CA Caprifoliaceae Valeriana sitchensis W north to AK

Cerastium fontanum ssp. vulgare (Cerastium Introduced throughout N America from Caryophyllaceae holosteoides) I Eurasia

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Honckenya peploides Sandy beaches of the PNW from OR north Caryophyllaceae ssp. major PNWcbic to AK; circumboreal

Rocky bluffs and gravel beaches of the coastal West from WA north to AK; Caryophyllaceae Sagina maxima CWica amphiberingian

Sagina maxima ssp. Rocky bluffs and gravel beaches of the Caryophyllaceae crassicaulis CWic" coastal West from CA north to AK

Rocky bluffs and gravel beaches of the Sagina maxima ssp. coastal West from WA north to AK; Caryophyllaceae maxima CWica" amphiberingian

Widespread in wet areas throughout N Caryophyllaceae Sagina procumbens NAcb America; circumboreal

Silene latifolia (Silene Introduced throughout N America from Caryophyllaceae alba) I Eurasia

Introduced throughout N America from Caryophyllaceae Silene vulgaris I Eurasia

Beaches and mudflats of the coastal West from CA north to AK; also beaches in NE Caryophyllaceae Spergularia canadensis NAic N America

Beaches and mudflats from BC north to Spergularia canadensis AK; also beaches of northeastern N Caryophyllaceae ssp. canadensis NNAic" America

Introduced throughout N America from Caryophyllaceae Spergularia rubra I Eurasia

Stellaria borealis ssp. Widespread throughout the West from CA Caryophyllaceae sitchana W north to AK

Widespread throughout wet areas of the Caryophyllaceae Stellaria calycanthus Wa West from CA north to AK; amphiberingian

Widespread in the west from CA north to Caryophyllaceae Stellaria crispa Wcb AK; circumboreal

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Mudflats and salt marshes of coastal BC Caryophyllaceae Stellaria humifusa CBCcbic and AK; circumboreal

Stellaria longipes ssp. Widespread throughout wet areas of N Caryophyllaceae longipes NAcb America; circumboreal

Introduced throughout N America from Caryophyllaceae Stellaria media I Eurasia

Widespread throughout wet areas of the Celastraceae Parnassia fimbriata W West from CA north to AK

Introduced to wet areas of the West from Convolvulaceae Calystegia sepium I eastern N America

Widespread throughout northern N Cornaceae Cornus canadensis NNAa America; amphiberingian

Widespread in wet areas across northern Cornus stolonifera N America and the coastal West from CA Cornaceae (Cornus sericea) NNAW north to AK

Wet areas of extreme NW BC north to AK; Cornaceae Cornus suecica CBCcb circumboreal

Widespread throughout the PNW from N Cornaceae Cornus unalaschkensis PNWa CA north to AK; amphiberingian

Infrequent in wet areas throughout the Crassulaceae Crassula aquatica Wcb West from CA north to AK; circumboreal

Rhodiola integrifolia ssp. Subalpine and alpine zones of the West Crassulaceae integrifolia Wa from CA north to AK; amphiberingian

Crassulaceae Sedum divergens CPNW Coastal PNW from OR north to SE AK

Chamaecyparis Cupressaceae nootkatensis CPNW Coastal PNW from CA north to SE AK

Widespread throughout N America; Cupressaceae Juniperus communis NAcb circumboreal

Juniperus communis var. Cupressaceae kelleyi PNW" PNW from OR north to AK

Huggins 35

Interior PNW and Great Basin; rare in N Cupressaceae Juniperus scopulorum PNW BC

Widespread throughout the PNW from N Cupressaceae Thuja plicata PNW CA north to SE AK

Frequent in alpine areas of coastal WA, Cyperaceae Carex anthoxanthea CPNW BC, and AK

Cyperaceae Carex aperta PNW Inland PNW; rare near the coast

Carex aquatilis ssp. Widespread throughout the West; 2 Cyperaceae aquatilis W subspecies

Cyperaceae Carex arcta PNW From NW CA north to BC

Cyperaceae Carex arctiformis CBC Coastal BC north to AK

Carex atrata ssp. Subalpine and alpine zones of the inland Cyperaceae atrosquama PNW PNW and Rocky Mountains

Cyperaceae Carex aurea W Widespread throughout the West

Cyperaceae Carex bebbii PNW Primarily inland PNW

Cyperaceae Carex bolanderi W Widespread throughout the West

Frequent from Sierra Nevadas north throughout PNW; circumboreal; also S. Cyperaceae Carex canescens PNWcbd America and Australia

Frequent from Sierra Nevadas north Carex canescens ssp. throughout PNW; circumboreal; also S. Cyperaceae canescens PNWcbd" America and Australia

Alpine areas of C BC; rare in immediate Cyperaceae Carex chordorrhiza PNW coastal OR and WA

Cyperaceae Carex circinata CPNW Coastal PNW from Puget Sound north

Cyperaceae Carex crawfordii PNW Throughout the PNW

Inland ranges from the Sierra Nevadas Cyperaceae Carex disperma PNW north

Widespread throughout wet areas of N Cyperaceae Carex echinata NAcb America; circumboreal

Huggins 36

Carex echinata ssp. echinata (Carex Widespread throughout N America; Cyperaceae phyllomanica) NAcb" circumboreal

Carex echinata ssp. Widespread throughout wet areas of the Cyperaceae phyllomanica W" West from CA north to AK

Rare in coastal salt marshes from N BC Cyperaceae Carex glareosa CBCcbic north; circumpolar

Carex glareosa var. Rare in coastal salt marshes from N BC Cyperaceae amphigena CBCcbic" north; circumpolar

Rare in extreme­coastal areas of BC and Cyperaceae Carex gmelinii CBCica AK; amphiberingian

Rare south of BC; scattered throughout Cyperaceae Carex gynocrates PNWa BC; amphiberingian

From the Sierra Nevadas and N Coast Cyperaceae Carex hoodii W Ranges north throughout the PNW

Cyperaceae Carex laeviculmis W Central CA north throughout the PNW

Cyperaceae Carex lasiocarpa PNW From N CA north throughout the PNW

Carex lenticularis (Carex Widespread throughout the West and Cyperaceae enanderi) NNAW northern N America; several subspecies

Carex lenticularis var. Widespread throughout the PNW from N Cyperaceae limnophila (Carex hindsii) PNW" CA north to AK

Carex lenticularis var. lipocarpa (Carex Widespread throughout the West from CA Cyperaceae kelloggii) W" north to AK

Carex leptalea ssp. Cyperaceae pacifica CBC Coastal BC and AK

Sierra Nevadas north to the Cascades and Cyperaceae Carex limosa W SW BC

Cyperaceae Carex livida PNW Infrequent and local south of coastal BC

Salt marshes and estuaries of the coastal Cyperaceae Carex lyngbyei CWica West from CA north to AK; amphiberingian

Huggins 37

Carex lyngbyei ssp. Salt marshes and estuaries of the coastal Cyperaceae cryptocarpa CWica" West from CA north to AK; amphiberingian

Rare and local in the PNW, more common Cyperaceae Carex macloviana PNW farther north

Sand beaches and dunes of the PNW; Cyperaceae Carex macrocephala CPNWa amphiberingian

Cyperaceae Carex macrochaeta PNWa Uncommon south of BC; amphiberingian

Frequent throughout WA and BC; Cyperaceae Carex magellanica NNAcb circumpolar

Carex magellanica ssp. irrigua (Carex Frequent throughout WA and BC; Cyperaceae paupercula) NNAcb" circumpolar

Wet areas of the West from CA north to Cyperaceae Carex mertensii Wa AK; amphiberingian

From the Sierra Nevadas north throughout Cyperaceae Carex nigricans W the Cascades and mountainous areas

Widespread throughout the near­coastal Cyperaceae Carex obnupta CW West

Wet areas of the PNW from NW WA north Cyperaceae Carex pauciflora NNAcb to AK; circumpolar

Cyperaceae Carex pluriflora CPNWa Near coastal PNW; amphiberingian

Cyperaceae Carex podocarpa PNWa Alpine areas of the PNW; amphiberingian

Carex pyrenaica ssp. Cyperaceae micropoda PNW Common in the PNW and Rockies

Peat bogs in montane areas of WA and Cyperaceae Carex rostrata NNAcb BC; circumpolar

Cyperaceae Carex saxatilis NNAcb Widespread in the PNW; circumpolar

Cyperaceae Carex saxatilis ssp. laxa NNAcb" Widespread in the PNW; circumpolar

Cyperaceae Carex sitchensis CPNW Widespread throughout the coastal PNW

Cyperaceae Carex spectabilis W Sierra Nevadas north throughout the PNW

Huggins 38

Common from central CA north throughout Cyperaceae Carex stipata W the PNW

NW WA north along near­coastal BC; Cyperaceae Carex stylosa CBCa amphiberingian

Sierra Nevadas and Bay Area north Cyperaceae Carex utriculata W throughout the PNW

Widespread in the West from the Sierra Cyperaceae Carex vesicaria W Nevadas north

NW CA north along the coast to AK; Cyperaceae Carex viridula CPNWa common in S BC; amphiberingian

Cyperaceae Eleocharis kamtschatica CBCa Near­coastal N BC; amphiberingian

Cyperaceae Eleocharis macrostachya W Widespread in wet areas of the West

Cyperaceae Eleocharis mamillata CPNW Near­coastal PNW

Common from central CA north throughout Cyperaceae Eleocharis obtusa CW the coastal PNW

Cyperaceae Eleocharis palustris W Widespread in wet areas of the West

Common from the Sierra Nevadas Cyperaceae Eleocharis quinqueflora W throughout the interior PNW

Cyperaceae Eriophorum angustifolium PNW Widespread in wet areas of the PNW

Widespread in wet areas across northern Cyperaceae Eriophorum chamissonis NNA N America

Eriophorum chamissonis Cyperaceae var. albidum PNW" Infrequent in N BC

Widespread in wet areas in BC; less Eriophorum chamissonis common in coastal OR and WA; also E Cyperaceae var. chamissonis NNA" Canada

Wet areas from the Sierra Nevadas north Cyperaceae Eriophorum gracile W throughout the PNW

Cyperaceae Rhynchospora alba W Wet areas from Central CA north to AK

Huggins 39

Schoenoplectus subterminalis (Scirpus Infrequent in wet areas from the Sierra Cyperaceae subterminalis) W Nevadas north

Schoenoplectus Cyperaceae tabernaemontani W Frequent in wet areas of the West

Cyperaceae Scirpus microcarpus W Widespread in wet areas of the West

Trichophorum cespitosum (Scirpus Wet areas throughout N America; Cyperaceae cespitosus) NAcb circumboreal

Gymnocarpium Widespread throughout the PNW from W Cystopteridaceae disjunctum PNWa OR north to AK; amphiberingian

Montane and subalpine zones throughout Cystopteridaceae Gymnocarpium dryopteris NNAcb northern N America; circumpolar

Dennstaedtiaceae Pteridium aquilinum C Cosmopolitan

Widespread in wet areas throughout Droseraceae Drosera anglica NNAcb northern N America; circumboreal

Widespread in wet areas throughout N Droseraceae Drosera rotundifolia NNAcb America; circumpolar

Dryopteris carthusiana (Dryopteris austriaca var. Widespread throughout northern N Dryopteridaceae dilatata) NNAcb America; circumboreal

Widespread throughout northern N Dryopteridaceae Dryopteris expansa NNAcb America; circumpolar

Dryopteridaceae Dryopteris filix­mas C Cosmopolitan

Infrequent in the (mostly) coastal PNW Dryopteridaceae Polystichum andersonii CPNW from OR north to AK

Infrequent in coastal BC and AK; Dryopteridaceae Polystichum braunii CBCcb circumpolar

Widespread in montane, subalpine, and alpine zones of the West from CA north to Dryopteridaceae Polystichum lonchitis Wcb AK; circumpolar

Huggins 40

Widespread throughout the West from CA Dryopteridaceae Polystichum munitum W north to SE AK

Widespread throughout the northern Equisetaceae Equisetum arvense NAcb hemisphere

Shallow water throughout northern N Equisetaceae Equisetum fluviatile NAcb America; circumboreal

Equisetaceae Equisetum hyemale NA Widespread throughout N America

Equisetum hyemale var. Equisetaceae affine NA" Widespread throughout N America

Widespread across montane and subalpine zones of northern N America; Equisetaceae Equisetum pratense NNAcb circumpolar

Widespread throughout the PNW and Equisetaceae Equisetum variegatum NNAcb northern N America; circumpolar

Andromeda polifolia Bogs throughout northern N America; Ericaceae (Kalmia polifolia) NNAcb circumboreal

Andromeda polifolia var. Bogs throughout northern N America; Ericaceae polifolia NNAcb" circumboreal

Infrequent and local in coastal BC from N Vancouver Island north to AK; Ericaceae Cassiope lycopodioides CBCa amphiberingian

Subalpine and alpine heath of the West Ericaceae Cassiope mertensiana W from CA north to SE AK

Cassiope mertensiana Subalpine and alpine heath of the West Ericaceae ssp. mertensiana W" from CA north to SE AK

Elliottia pyroliflorus (Cladothamnus Montane, subalpine, and alpine zones of Ericaceae pyroliflorus) CPNW the coastal PNW from OR north to AK

Widespread throughout northern N Ericaceae Empetrum nigrum NNAcb America; circumpolar

Huggins 41

Empetrum nigrum ssp. Widespread throughout northern N Ericaceae nigrum NNAcb" America; circumpolar

Widespread throughout the coastal PNW Ericaceae Gaultheria shallon CPNW from N CA north to AK

Subalpine and alpine heath of the coastal Harrimanella stelleriana PNW from WA north to AK; Ericaceae (Cassiope stelleriana) CPNWa amphiberingian

Ericaceae Kalmia microphylla W Bogs of the West from CA north to AK

Kalmia microphylla var. occidentalis (Kalmia Ericaceae occidentalis) W" Bogs of the West from CA north to AK

Kalmia procumbens Bogs and alpine heath throughout Ericaceae (Loiseleuria procumbens) NNAcb northern N America; circumpolar

Widespread throughout the PNW from N Ericaceae Menziesia ferruginea PNW CA north to AK

Menziesia ferruginea ssp. Widespread throughout the PNW from N Ericaceae ferruginea PNW" CA north to AK

Widespread throughout northern N Ericaceae Moneses uniflora NNAcb America; circumboreal

Monotropa hypopithys Widespread throughout N America; Ericaceae (Hypopitys monotropa) NNAcb circumboreal

Orthilia secunda (Pyrola Widespread throughout northern N Ericaceae secunda) NNAcb America; circumboreal

Orthilia secunda var. Widespread throughout northern N Ericaceae secunda NNAcb" America; circumboreal

Phyllodoce aleutica ssp. Ericaceae glanduliflora CBCa AK; amphiberingian

Montane, subalpine, and alpine zones of Ericaceae Phyllodoce empetriformis PNW the PNW from N CA north to AK

Montane, subalpine, and alpine zones of Ericaceae Phyllodoce glanduliflora PNW the PNW from OR north to AK

Huggins 42

Pyrola asarifolia ssp. Widespread across northern N America; Ericaceae asarifolia NNAa amphiberingian

Rhododendron groenlandicum (Ledum Ericaceae groenlandicum) NNA Bogs throughout northern N America

Ericaceae Vaccinium alaskaense CPNW Coastal PNW from NW OR north to AK

Widespread throughout northern N Ericaceae Vaccinium caespitosum NNA America

Widespread in montane and subalpine zones of the PNW from N CA north to N Vaccinium BC; locally infrequent on the north coast of Ericaceae membranaceum PNW BC

Widespread throughout the PNW from OR north to AK; amphiberingian; disjunct to E Ericaceae Vaccinium ovalifolium PNWad Canada

Vaccinium oxycoccos Bogs throughout northern N America; Ericaceae (Oxycoccos oxycoccos) NNAcb circumboreal

Widespread throughout the coastal PNW Ericaceae Vaccinium parvifolium CPNW from N CA north to SE AK

Bogs and alpine heath throughout Ericaceae Vaccinium uliginosum NNAcb northern N America; circumboreal

Bogs and alpine heath throughout Vaccinium uliginosum northern N America; mostly coastal BC; Ericaceae ssp. occidentale NNAcb" circumboreal

Bogs and alpine heath throughout Vaccinium uliginosum northern N America; mostly N BC; Ericaceae ssp. pubescens NNAcb" circumboreal

Bogs and alpine heath throughout Ericaceae Vaccinium vitis­idaea NNAcb northern N America; circumboreal

Vaccinium vitis­idaea Bogs and alpine heath throughout Ericaceae ssp. minus NNAcb" northern N America; circumboreal

Huggins 43

Introduced throughout N America from Euphorbiaceae Euphorbia esula I Eurasia

Introduced throughout the coastal PNW Fabaceae Cytisus scoparius I from Eurasia

Beaches of the PNW from N CA north to Fabaceae Lathyrus japonicus CPNWcbicd AK; circumboreal; also S America

Wet areas of the coastal PNW from N CA Fabaceae Lathyrus palustris CPNW north to AK

Coastal BC and AK; also Canadian Fabaceae Lupinus nootkatensis CBC Rockies

Lupinus nootkatensis var. Fabaceae nootkatensis CBC" Coastal BC and AK

Widespread throughout the West from CA north to AK and across northern N Fabaceae Lupinus polyphyllus NNAW America

Widespread throughout the West from CA Lupinus polyphyllus ssp. north to AK and across northern N Fabaceae polyphyllus NNAW" America

Introduced throughout N America from Fabaceae Melilotus albus I Eurasia

Introduced throughout N America from Fabaceae Melilotus officinalis I Eurasia

Introduced throughout N America from Fabaceae Trifolium hybridum I Eurasia

Introduced throughout N America from Fabaceae Trifolium pratense I Eurasia

Infrequent in wet areas of the West from Fabaceae Trifolium wormskioldii W CA north to NW BC

Introduced throughout N America from Fabaceae Vicia cracca I Eurasia

Huggins 44

Vicia nigricans ssp. Widespread in the coastal West from CA Fabaceae gigantea CW north to AK

Wet areas of the coastal PNW from WA Gentianaceae Gentiana douglasiana CPNW north to AK

Wet areas of subalpine and alpine zones of coastal BC from N Vancouver Island Gentianaceae Gentiana platypetala CBC north to SE AK

Wet areas of the coastal PNW from N CA Gentianaceae Gentiana sceptrum CPNW north to NW BC

Wet areas of the coastal West from CA Grossulariaceae Ribes bracteosum CW north to AK

Widespread throughout the coastal West Grossulariaceae Ribes divaricatum CW from CA north to coastal BC

Widespread throughout the interior West Grossulariaceae Ribes hudsonianum NNAW and across northern N America

Widespread throughout wet montane and subalpine zones of the West from CA north to AK and across northern N Grossulariaceae Ribes lacustre NNAW America

Widespread throughout the PNW from N Grossulariaceae Ribes laxiflorum PNW CA north to AK

Widespread throughout wet areas of the Haloragaceae Myriophyllum sibiricum Wcb West from CA north to AK; circumboreal

Hypericaceae Hypericum anagalloides W Wet areas of the west from CA north to AK

Introduced throughout N America from Iridaceae Iris pseudacorus I Eurasia

Sisyrinchium Introduced throughout the West from Iridaceae angustifolium I eastern N America

From SW WA north to Alaska along the Iridaceae Sisyrinchium littorale CPNWic immediate coast

Huggins 45

Wet areas of the coastal PNW from W WA Isoetaceae Isoetes maritima CPNWa north to AK; amphiberingian

Wet areas throughout the West from CA Isoetes tenella (Isoetes north to AK and across northern N Isoetaceae echinospora) NNAWcb America; circumboreal

Common in wet areas of the PNW; less common in CA; disjunct in eastern N Juncaceae Juncus acuminatus CPNWd America

Infrequent in wet areas across northern N Juncaceae Juncus alpinoarticulatus NNAcb America; circumboreal

Wet areas throughout N America; Juncaceae Juncus arcticus NAcbd circumboreal; also S America

Juncus arcticus ssp. Juncaceae alaskanus CBC" Infrequent in wet areas of BC north to AK

Juncus arcticus ssp. Infrequent in wet areas of BC north to AK; Juncaceae sitchensis CBCa" amphiberingian

Widespread in wet areas of the West from CA north to AK and across northern N Juncaceae Juncus articulatus NNAWcb America; circumboreal

Widespread in wet areas of the N Juncaceae Juncus balticus NAcb America; circumboreal

Widespread in wet areas of the N Juncaceae Juncus balticus ssp. ater NAcb" America; circumboreal

Sand dunes of the immediate coastal Juncaceae Juncus breweri CWic West from CA north to NW BC

Juncaceae Juncus bufonius C Cosmopolitan

Juncaceae Juncus bulbosus I Introduced throughout the coastal PNW

Introduced to wet areas of the coastal Juncaceae Juncus conglomeratus I PNW

Wet areas from CA north throughout the Juncaceae Juncus covillei CW coastal PNW

Huggins 46

Juncaceae Juncus drummondii W Common in wet areas of the alpine West

Juncaceae Juncus effusus C Cosmopolitan plant of wet areas

Widespread and common in wet areas of Juncus effusus ssp. the West from CA north to AK; also Juncaceae effusus Wd" Eurasia

Juncus effusus ssp. Widespread in the coastal West from CA Juncaceae pacificus CW" north to AK

Widespread and common in wet areas of the West from CA north to AK and across Juncaceae Juncus ensifolius NNAWa northern N America; amphiberingian

Frequent in wet areas of the near­coastal West from CA north to AK; Juncaceae Juncus falcatus CWad amphiberingian; also Australia

Frequent in wet areas of the near­coastal Juncus falcatus ssp. West from CA north to AK; Juncaceae sitchensis CWad" amphiberingian; also Australia

Infrequent in wet areas across northern N Juncaceae Juncus filiformis NNAcb America; circumboreal

Rare and local to wet areas of Haida Gwaii Juncaceae Juncus haenkei CBCa and coastal Alaska; amphiberingian

Wet areas in montanze zones from the Sierra Nevadas north throughout the Juncaceae Juncus mertensianus Wa PNW; amphiberingian

Bogs of northern Vancouver Island and N BC and across northern N America; Juncaceae Juncus stygius NNAcb circumpolar

Bogs of northern Vancouver Island and N Juncus stygius ssp. BC and across northern N America; Juncaceae americanus NNAcb" circumpolar

Wet areas of the near­coast from Marin Juncaceae Juncus supiniformis CW County CA north to AK

Huggins 47

Common throughout wet areas of N Juncaceae Juncus tenuis NA America

Juncaceae Luzula arcuata PNWa Infrequent in alpine BC; amphiberingian

Juncaceae Luzula campestris I Introduced throughout the PNW

Widespread throughout the West; 2 Juncaceae Luzula comosa W subspecies

Luzula comosa var. Juncaceae comosa W" Widespread throughout the West

Common in the PNW, but rare and local in Juncaceae Luzula multiflora C CA; cosmopolitan

Luzula multiflora ssp. Common in the PNW, but rare and local in Juncaceae frigida C" CA; cosmopolitan

Luzula multiflora ssp. Common in the PNW, but rare and local in Juncaceae multiflora C" CA; cosmopolitan

Frequent in alpine areas from the Sierra Nevadas north to Alaska throughout the Juncaceae Luzula parviflora NNAWcb PNW

Frequent in alpine areas from the Sierra Nevadas north to Alaska throughout the Luzula parviflora ssp. PNW and across northern N America; Juncaceae fastigiata NNAWcb" circumboreal

Frequent in alpine areas of extreme NW Juncaceae Luzula piperi PNWa CA and the PNW; amphiberingian

Infrequent in wet areas of the West from Juncaginaceae Lilaea scilloides Wd CA north to NW BC; also S. America

Tidal marshes and alkali flats of the coastal West from CA north to NW BC; Juncaginaceae Triglochin concinna CWic also S. America

Tidal marshes and alkali flats of the Triglochin concinna var. coastal West from CA north to NW BC; Juncaginaceae concinna CWic" also S. America

Huggins 48

Juncaginaceae Triglochin maritima C Cosmopolitan plant of wet areas

Cosmopolitan plant of wet areas; Juncaginaceae Triglochin palustris C infrequent along the coast

Widespread in wet areas throughout N Lamiaceae Lycopus uniflorus NAa America; amphiberingian

Widespread throughout N America; Lamiaceae Prunella vulgaris NAcb circumboreal

Prunella vulgaris ssp. Lamiaceae lanceolata NA" Widespread throughout N America

Stachys chamissonis var. cooleyi (Stachys Wet areas of the coastal PNW from W OR Lamiaceae cooleyae) CPNW north to NW BC

Stachys mexicana Widespread in wet areas of the coastal Lamiaceae (Stachys emersonii) CW West from CA north to SE AK

Wet areas throughout northern N America; Lentibulariaceae Pinguicula vulgaris NNAcb circumboreal

Pinguicula vulgaris ssp. macroceras (Pinguicula Wet areas throughout the coastal West Lentibulariaceae macroceras) CWa" from CA north to AK; amphiberingian

Pinguicula vulgaris ssp. Wet areas throughout northern N America; Lentibulariaceae vulgaris NNAcb" circumboreal

Wet areas throughout northern N America; Lentibulariaceae Utricularia intermedia NNAcb circumboreal

Utricularia macrorhiza Wet areas throughout N America; Lentibulariaceae (Utricularia vulgaris) NNAa amphiberingian

Wet areas throughout northern N America; Lentibulariaceae Utricularia minor NNAcb circumboreal

Infrequent in wet montane areas of the Lentibulariaceae Utricularia ochroleuca PNWcb PNW and E Canada; circumboreal

Widespread throughout the PNW from N Liliaceae Clintonia uniflora PNW CA north to AK

Huggins 49

Wet areas of coastal BC and AK; rare and Liliaceae Fritillaria camschatcensis CPNW local in coastal OR and WA

Fritillaria camschatcensis Wet areas of coastal BC and AK; rare and Liliaceae ssp. camschatcensis CPNW" local in coastal OR and WA

Widespread throughout N America; Liliaceae Streptopus amplexifolius NAcb circumboreal

Streptopus lanceolatus (Streptopus roseus var. Widespread throughout the coastal PNW Liliaceae curvipes) CPNW from OR north to SE AK

Montane and subalpine zones of the Streptopus streptopoides coastal PNW from WA north to AK; Liliaceae var. brevipes CPNWa amphiberingian

Widespread throughout N America; Linnaeaceae Linnaea borealis NAcb circumboreal

Linnaea borealis ssp. Linnaeaceae longiflora (americana) NA" Widespread throughout N America

Diphasiastrum alpinum Subalpine and alpine zones throughout Lycopodiaceae (Lycopodium alpinum) NNAcb northern N America; circumpolar

Diphasiastrum complanatum (Lycopodium Widespread throughout northern N Lycopodiaceae complanatum) NNAcbd America; circumpolar; S America

Huperzia chinensis Wet areas of the PNW from OR north to Lycopodiaceae (Huperzia miyoshiana) PNWa AK; amphiberingian

Subalpine and alpine zones of the PNW Lycopodiaceae Huperzia haleakalae PNWa from OR north to AK; amphiberingian?

Wet areas of the PNW from OR north to Lycopodiaceae Huperzia occidentalis PNW AK

Lycopodiella inundata Wet areas across northern N America; Lycopodiaceae (Lycopodium inundatum) NNAcb circumboreal

Huggins 50

Widespread throughout northern N Lycopodiaceae Lycopodium annotinum NNAcb America; circumpolar

Lycopodiaceae Lycopodium clavatum C Cosmopolitan

Lycopodium clavatum Lycopodiaceae ssp. clavatum C" Cosmopolitan

Lycopodium clavatum Lycopodiaceae var. monostachyon C" Cosmopolitan

Lycopodium Widespread throughout northern N Lycopodiaceae dendroideum NNAa America; amphiberingian

Widespread throughout northern N Lycopodiaceae Lycopodium lagopus NNAcb America; circumboreal

Lycopodium sitchense (Diphasiastrum Subalpine and alpine zones throughout Lycopodiaceae sitchense) NNAa northern N America; amphiberingian

Veratrum viride var. eschscholzianum Widespread in wet areas of the West from Melanthiaceae (Veratrum eschscholtzii) Wd CA north to AK; disjunct in east N America

Wet areas throughout the West and Menyanthaceae Menyanthes trifoliata NNAWcb across northern N America; circumboreal

Nephrophyllidium crista­galli (Fauria Wet areas of the coastal PNW from WA Menyanthaceae crista­galli) CPNWa north to AK; amphiberingian

Wet areas throughout the West from CA Montiaceae Claytonia lanceolata W north to NW BC

Claytonia sibirica ssp. Wet areas of the (mostly) coastal West Montiaceae sibirica CWa from CA north to AK; amphiberingian

Wet areas throughout the West and Montiaceae Montia fontana NNAWcb across northern N America; circumboreal

Wet areas throughout the West from CA Montiaceae Montia parviflora W north to SE AK

Huggins 51

Montia parviflora var. Wet areas throughout the West from CA Montiaceae flagellaris W" north to SE AK

Montia parvifolia var. Wet areas throughout the West from CA Montiaceae parvifolia W" north to SE AK

Wet areas throughout northern N America; Myricaceae Myrica gale NNAcb circumboreal

Nuphar polysepala (Nuphar lutea ssp. Wet areas throughout the West from CA Nymphaeaceae polysepala) W north to AK

Rare in wet areas of the PNW from WA Nymphaeaceae Nymphaea tetragona PNW north to AK

Widespread throughout N America; Onagraceae Circaea alpina NAcb circumboreal

Widespread throughout N America; Onagraceae Circaea alpina ssp. alpina NAcb" circumboreal

Epilobium Widespread throughout montane and Onagraceae anagallidifolium NAcb subalpine N America; circumboreal

Epilobium angustifolium (Chamerion Widespread throughout N America; Onagraceae angustifolium) NAcb circumboreal

Widespread throughout N America; Onagraceae Epilobium ciliatum NAa amphiberingian

Epilobium ciliatum ssp. Widespread throughout N America; Onagraceae ciliatum NAa" amphiberingian

Epilobium ciliatum ssp. glandulosum (Epilobium Widespread throughout N America; Onagraceae glandulosum) NAa" amphiberingian

Epilobium ciliatum ssp. Widespread throughout N America; Onagraceae watsonii NAa" amphiberingian

Epilobium hornemannii Wet areas throughout the West and Onagraceae ssp. hornemannii NNAWcb across northern N America; circumboreal

Huggins 52

Wet montane and subalpine zones throughout the West and across northern Onagraceae Epilobium latifolium NNAWcb N America; circumboreal

Wet areas across northern N America; locally abundant in the Skeena River Onocleaceae Matteuccia struthiopteris NNAcb valley; circumboreal

Widespread throughout the West and Orchidaceae Calypso bulbosa NNAWcb across northern N America; circumboreal

Widespread throughout N America; also C Orchidaceae Corallorhiza maculata NA America

Corallorhiza mertensiana (Corallorhiza maculata Widespread throughout the West from CA Orchidaceae ssp. mertensiana) W north to AK

Infrequent in wet areas across northern N Orchidaceae Malaxis brachypoda NNAad America; disjunct in CA; amphiberingian

Malaxis paludosa Infrequent in wet areas across northern N Orchidaceae (Hammarbya paludosa) NNAcb America; circumboreal

Neottia caurina (Listera Wet areas in the montane and subalpine Orchidaceae caurina) W zones of the West from CA north to AK

Neottia cordata (Listera Wet areas throughout N America; Orchidaceae cordata) NAcb circumboreal

Wet areas throughout the West and Orchidaceae Piperia unalascensis NNAW across northern N America

Platanthera aquilonis (Platanthera / Habenaria Widespread in wet montane and subalpine Orchidaceae hyperborea) NNA areas across northern N America

Wet areas of the coastal PNW from W WA Orchidaceae Platanthera chorisiana CPNWa north to AK; amphiberingian

Wet areas throughout the West and Orchidaceae Platanthera dilatata NNAW across northern N America

Huggins 53

Platanthera dilatata var. Wet areas of the coastal PNW from N CA Orchidaceae leucostachys CPNW north to AK

Orchidaceae Platanthera orbiculata NA Wet montane areas throughout N America

Platanthera stricta Wet montane areas of the West from CA Orchidaceae (Platanthera saccata) W north to AK

Widespread throughout the West and across northern N America; also the Orchidaceae Spiranthes romanzoffiana NNAWd British Isles

Widespread throughout the West and Spiranthes romanzoffiana across northern N America; also the Orchidaceae var. romanzoffiana NNAWd" British Isles

Widespread throughout the West from CA Orobanchaceae Castilleja miniata Wd north to AK; also NE N America

Castilleja miniata var. Widespread throughout the West from CA Orobanchaceae miniata Wd" north to AK; also NE N America

Wet areas of subalpine and alpine zones Orobanchaceae Castilleja parviflora W of the West from CA north to AK

Castilleja parviflora var. Wet areas of subalpine and alpine zones Orobanchaceae parviflora W" of the West from CA north to AK

Introduced throughout the coastal PNW Orobanchaceae Euphrasia nemorosa I from Eurasia

Orobanchaceae Melampyrum lineare NNA Infrequent across northern N America

Pedicularis Subalpine and alpine zones of the coastal Orobanchaceae ornithorhyncha CPNW PNW from WA north to AK

Rare in wet montane and subalpine zones Orobanchaceae Pedicularis parviflora NNAcb across northern N America; circumboreal

Pedicularis parviflora var. Rare in wet montane and subalpine zones Orobanchaceae parviflora NNAcb" across northern N America; circumboreal

Wet subalpine and alpine zones of N BC Orobanchaceae Pedicularis sudetica CBCcb north to AK; circumpolar

Huggins 54

Erythranthe guttata Widespread in wet areas of the West from Phrymaceae (Erythranthe microphylla) Wd CA north to AK; disjunct in east N America

Widespread throughout the coastal PNW from N CA north to SE AK; absent from Pinaceae Abies amabilis CPNW Haida Gwaii

Picea engelmannii × Occurs where the ranges of these two Pinaceae Picea glauca PNW" species meet in central BC

Widespread throughout the coastal PNW Pinaceae Picea sitchensis CPNW from N CA north to SE AK

Widespread throughout the West from CA Pinaceae Pinus contorta W north to AK

Pinus contorta var. Widespread throughout the West from CA Pinaceae contorta W" north to AK

Widespread throughout the coastal PNW Pinaceae Tsuga heterophylla CPNW from N CA north to SE AK

Montane and subalpine zones of the Pinaceae Tsuga mertensiana CW coastal West from CA north to SE AK

Callitriche heterophylla Wet areas of the coastal West from CA Plantaginaceae ssp. bolanderi CW north to AK

Callitriche palustris Wet areas throughout N America; Plantaginaceae (Callitriche verna) NAcb circumboreal

Introduced throughout N America from Plantaginaceae Digitalis purpurea I Eurasia

Wet areas of the PNW from W WA north Plantaginaceae Hippuris montana PNWa to AK; amphiberingian

Montane, subalpine, and alpine zones of Plantaginaceae Penstemon davidsonii CPNW the coastal PNW from N CA north to BC

Wet areas of the coastal PNW from W OR Plantaginaceae Penstemon serrulatus CPNW north to NW BC

Wet areas of the coastal PNW from W OR Plantaginaceae Plantago macrocarpa CPNWa north to AK; amphiberingian

Huggins 55

Introduced throughout N America from Plantaginaceae Plantago major I Eurasia

Plantaginaceae Plantago maritima Cic Coastal ­ Cosmopolitan

Plantago maritima ssp. Beaches and salt marshes of the Pacific Plantaginaceae juncoides CWic" coast from S America north to AK

Veronica Introduced throughout N America from Plantaginaceae anagallis­aquatica I Eurasia

Veronica beccabunga ssp. americana (Veronica Plantaginaceae americana) NA Wet areas throughout N America

Veronica beccabunga Introduced throughout N America from Plantaginaceae ssp. beccabunga I Eurasia

Introduced throughout N America from Poaceae Aegilops tauschii I Eurasia

Introduced throughout N America from Poaceae Agrostis capillaris I Eurasia

Widespread throughout the West from CA north to AK; amphiberingian; also S Poaceae Agrostis exarata Wad America

Introduced throughout N America from Poaceae Agrostis gigantea I Eurasia

Widespread across northern N America; Poaceae Agrostis mertensii NNAcb circumboreal

Wet areas of the PNW from N CA north to Poaceae Agrostis oregonensis PNW NW BC

Rocky slopes and dunes of the lowland Poaceae Agrostis pallens W West from CA north to NW BC

Agrostis scabra (Agrostis Widespread throughout N America; Poaceae hiemalis) NAa amphiberingian; also Greenland

Introduced throughout N America from Poaceae Agrostis stolonifera I Eurasia

Huggins 56

Agrostis stolonifera var. Introduced throughout N America from Poaceae majus I Eurasia

Subalpine and alpine zones of the West Poaceae Agrostis variabilis W from CA north to N BC

Introduced throughout N America from Poaceae Alopecurus geniculatus I Eurasia

Wet areas of immediate coastal NW BC Arctopoa eminens (Poa north to AK; amphiberingian; disjunct to Poaceae eminens) CBCad NE N America

Widespread throughout the West from CA Poaceae Bromus carinatus W north to AK

Bromus carinatus var. Widespread throughout the West from CA Poaceae carinatus W" north to AK

Bromus carinatus var. Widespread throughout the West from CA Poaceae marginatus W" north to AK

Poaceae Bromus pacificus CPNW Coastal PNW from W OR north to SE AK

Bromus sitchensis Poaceae (Bromus aleutensis) CPNW Coastal PNW from W OR north to AK

Bromus squarrosus var. Introduced throughout N America from Poaceae squarrosus I Eurasia

Introduced throughout N America from Poaceae Bromus tectorum I Eurasia

Widespread throughout the West from CA Poaceae Bromus vulgaris W north to NW BC

Calamagrostis Widespread throughout N America; Poaceae canadensis NAcb circumboreal

Calamagrostis canadensis var. Poaceae canadensis NA" Widespread throughout N America

Huggins 57

Calamagrostis canadensis var. Widespread throughout N America; Poaceae langsdorfii NAcb" circumboreal

Widespread in the near­coastal West from Poaceae Calamagrostis nutkaensis CW CA north to AK

Widespread throughout the West from CA Calamagrostis stricta ssp. north to AK and across northern N Poaceae inexpansa NNAWa America; amphiberingian

Widespread throughout the West from CA north to AK and across northern N Poaceae Cinna latifolia NNAWcb America; circumboreal

Introduced throughout N America from Poaceae Dactylis glomerata I Eurasia

Widespread throughout the West from CA north to AK and across northern N Poaceae Danthonia intermedia NNAW America

Poaceae Danthonia spicata NA Widespread throughout N America

Widespread in wet areas throughout N Poaceae Deschampsia cespitosa NAcb America; circumboreal

Deschampsia cespitosa var. beringensis (Deschampsia Wet areas of the coastal West from CA Poaceae beringensis) CWa north to AK; amphiberingian

Dichanthelium acuminatum ssp. Poaceae fasciculatum NA Widespread throughout N America

Poaceae Elymus glaucus NA Widespread throughout N America

Elymus glaucus ssp. Poaceae glaucus NA" Widespread throughout N America

Poaceae Elymus hirsutus CPNW Coastal PNW from W OR north to SE AK

Huggins 58

Elymus repens (Elytrigia Introduced throughout N America from Poaceae pungens) I Eurasia

Festuca filiformis Introduced throughout N America from Poaceae (Festuca tenuifolia) I Eurasia

Widespread in wet areas throughout N Poaceae Festuca rubra NAcb America; circumboreal

Festuca rubra ssp. Wet areas of the coastal West from CA Poaceae pruinosa CWa north to AK; amphiberingian

Widespread throughout the West from CA Poaceae Festuca subulata W north to SE AK

Wet areas throughout the West from CA north to AK and across northern N Poaceae Glyceria borealis NNAW America

Wet areas throughout the West from CA north to AK and across northern N Poaceae Glyceria grandis NNAW America

Poaceae Glyceria striata NA Wet areas throughout N America

Wet areas of the West from CA north to Poaceae Glyceria x occidentalis W NW BC

Widespread throughout the West from CA north to AK and across northern N Poaceae Hierochloe hirta NNAWcb America; circumboreal

Introduced throughout N America from Poaceae Holcus lanatus I Eurasia

Hordeum Poaceae brachyantherum NA Widespread throughout N America

Hordeum jubatum ssp. Widespread throughout N America; Poaceae jubatum NAcbd circumboreal; also S America

Leymus mollis (Elymus mollis) (Leymus Beaches and dunes of the coastal West Poaceae arenarius) CWcbic from CA north to AK; circumboreal

Huggins 59

Beaches and dunes of the coastal West Poaceae Leymus mollis ssp. mollis CWcbic" from CA north to AK; circumboreal

Introduced throughout N America from Poaceae Phalaris arundinacea I Eurasia

Widespread throughout N America; Poaceae Phleum alpinum NAcbd circumboreal; also S America

Introduced throughout N America from Poaceae Phleum pratense I Eurasia

Introduced throughout N America from Poaceae Poa annua I Eurasia

Wet montane and subalpine zones across Poaceae Poa arctica ssp. arctica NNAcb northern N America; circumboreal

Introduced throughout N America from Poaceae Poa compressa I Eurasia

Montane and subalpine zones of the West Poaceae Poa cusickii ssp. epilis W from CA north to AK

Infrequent throughout the PNW from W Poaceae Poa laxiflora PNW OR north to SE AK

Beaches and dunes of the coastal West Poaceae Poa macrantha CWic from CA north to AK

Poa nemoralis (Poa Poaceae interior) NA Montane zones throughout N America

Widespread in wet areas throughout N Poaceae Poa palustris NAcb America; circumboreal

Alpine zones of the PNW from WA north to Poaceae Poa paucispicula PNWa AK; amphiberingian

Poaceae Poa pratensis NAcb Northern hemisphere

Poa pratensis ssp. Introduced throughout N America from Poaceae irrigata I Eurasia

Poa pratensis ssp. Introduced throughout N America from Poaceae pratensis I Eurasia

Huggins 60

Poaceae Poa stenantha PNW PNW from OR north to AK

Podagrostis aequalis Wet areas of the coastal PNW from W OR Poaceae (Agrostis aequivalvis) CPNW north to SE AK

Podagrostis humilis Wet montane and subalpine zones Poaceae (Agrostis thurberiana) W throughout the West from CA north to AK

Beaches and wet areas of the coastal West from CA north to AK; disjunct at the Poaceae Puccinellia nutkaensis CWicd mouth of the St. Lawrence

Wet alkaline areas throughout the West from CA north to AK and across northern Poaceae Puccinellia nuttalliana NNAW N America

Beaches and wet areas of the coastal Poaceae Puccinellia pumila CWic West from CA north to AK

Schedonorus Introduced throughout N America from Poaceae arundinaceus I Eurasia

Introduced throughout N America from Poaceae Thinopyrum intermedium I Eurasia

Torreyochloa pauciflora Wet areas throughout the West from CA Poaceae (Puccinellia pauciflora) W north to AK

Widespread throughout the West from CA Poaceae Trisetum canescens W north to AK

Infrequent throughout the West from CA Poaceae Trisetum cernuum W north to AK

Widespread throughout N America; Poaceae Trisetum spicatum NAcbd circumboreal; also S America

Wet montane and subalpine zones of the West from CA north to AK and across northern N America; circumboreal; also S Poaceae Vahlodea atropurpurea NNAWcbd America

Huggins 61

Polemonium pulcherrimum ssp. Montane, subalpine, and alpine zones of Polemoniaceae lindleyi W the West from CA north to AK

Bistorta vivipara Montane subalpine zones across northern Polygonaceae (Polygonum viviparum) NNAcb N America; circumboreal

Fallopia convolvulus Introduced throughout N America from Polygonaceae (Polygonum convolvulus) I Eurasia

Persicaria wallichii (Polygonum Introduced throughout N America from Polygonaceae polystachyum) I Eurasia

Introduced throughout N America from Polygonaceae Rumex acetosella I Eurasia

Rumex aquaticus var. fenestratus (Rumex fenestratus) (Rumex Wet areas of the West from CA north to Polygonaceae occidentalis) NNAW AK and across northern N America

Rumex salicifolius Wet areas of the West from CA north to Polygonaceae (Rumex transitorius) W AK

Widespread throughout the coastal West Polypodiaceae Polypodium glycyrrhiza CW from CA north to SE AK

Widespread throughout the West from CA Polypodiaceae Polypodium hesperium W north to AK

Wet areas of the West from CA north to AK and across northern N America; Potamogetonaceae Potamogeton alpinus NNAWcb circumboreal

Wet areas throughout N America; disjunct Potamogetonaceae Potamogeton epihydrus NAd in parts of Europe

Wet areas of the West from CA north to AK and across northern N America; Potamogetonaceae Potamogeton gramineus NNAWcb circumboreal

Huggins 62

Wet areas of the West from CA north to AK and across northern N America; Potamogetonaceae Potamogeton natans NNAWcb circumboreal

Potamogeton pusillus Wet areas throughout N America; Potamogetonaceae ssp. tenuissimus NAcbd circumboreal; also Africa

Wet areas of the coastal PNW from N CA Primulaceae Dodecatheon jeffreyi CPNW north to AK

Dodecatheon jeffreyi ssp. Wet areas of the coastal PNW from N CA Primulaceae jeffreyi CPNW" north to AK

Dodecatheon pulchellum Widespread in wet areas of the West from Primulaceae ssp. pulchellum W CA north to AK

Dodecatheon pulchellum Widespread in wet areas of the West from Primulaceae var. macrocarpum W" CA north to AK

Widespread in wet areas throughout N Primulaceae Glaux maritima NAcb America; circumboreal

Trientalis europaea ssp. arctica (Lysimachia Wet areas of the coastal West from CA Primulaceae europaea) CWd north to AK; also Eurasia

Adiantum aleuticum Widespread throughout the West from CA (Adiantum pedatum ssp. north to AK; amphiberingian; disjunct to Pteridaceae aleuticum) Wad NE N America

Widespread throughout rocky areas of the Cryptogramma West from CA north to AK and across Pteridaceae acrostichoides NNAWa northern N America; amphiberingian

Wet rocks across northern N America; Pteridaceae Cryptogramma stelleri NNAcb circumboreal

Aconitum delphinifolium Montane and subalpine zones of N BC Ranunculaceae ssp. chamissonianum CBCa north to AK; amphiberingian

N BC north to AK; infrequent in coastal BC Ranunculaceae Anemone narcissiflora CBC from N Vancouver island north

Huggins 63

Wet montane and subalpine zones across Ranunculaceae Anemone parviflora NNAcb northern N America; circumboreal

Widespread in wet areas of the West from Ranunculaceae Aquilegia formosa W CA north to AK

Caltha leptosepala Widespread in wet areas of the West from Ranunculaceae (Caltha biflora) W CA north to AK

Caltha leptosepala ssp. Widespread in wet areas of the West from Ranunculaceae howellii W" CA north to AK

Caltha leptosepala var. Widespread in wet areas of the West from Ranunculaceae biflora W" CA north to AK

Caltha palustris ssp. Infrequent in wet areas across northern N Ranunculaceae asarifolia NNAcb America; circumboreal

Wet areas of the coastal PNW from NW Ranunculaceae Coptis aspleniifolia CPNW WA north to AK

Wet areas of across northern N America; Ranunculaceae Coptis trifolia NNAa amphiberingian

Wet areas of subalpine and alpine zones Kumlienia cooleyae of the coastal PNW from W WA north to Ranunculaceae (Ranunculus cooleyae) CPNW AK

Introduced throughout N America from Ranunculaceae Ranunculus acris I Eurasia

Wet areas throughout N America; Ranunculaceae Ranunculus aquatilis NAcbd circumboreal; also S America

Wet areas throughout N America; Ranunculaceae Ranunculus cymbalaria NAcb circumboreal

Ranunculus eschscholtzii Montane and subalpine zones throughout Ranunculaceae var. eschscholtzii W the West from CA north to AK

Wet areas throughout the West from CA north to AK and across northern N Ranunculaceae Ranunculus flammula NNAWcb America; circumboreal

Huggins 64

Ranunculus flammula Wet areas throughout the West from CA var. reptans (Ranunculus north to AK and across northern N Ranunculaceae reptans) NNAWcb" America; circumboreal

Wet areas throughout the West from CA north to AK and across northern N Ranunculaceae Ranunculus macounii NNAW America

Wet areas throughout the West from CA Ranunculus macounii north to AK and across northern N Ranunculaceae var. macounii NNAW" America

Widespread throughout the West from CA Ranunculaceae Ranunculus occidentalis W north to AK

Ranunculus Wet areas throughout the West from CA Ranunculaceae ornithorhyncha W north to SE AK

Ranunculus ornithorhyncha var. Wet areas throughout the West from CA Ranunculaceae platyphyllus W" north to SE AK; more frequent inland

Ranunculus uncinatus Wet areas of the West from CA north to Ranunculaceae (Ranunculus bongardii) W AK

Widespread in wet areas throughout N Ranunculaceae Trautvetteria caroliniensis NAa America; amphiberingian

Trollius albiflorus (Trollius Subalpine and alpine zones of the PNW Ranunculaceae laxus) PNW from WA north to BC

Widespread throughout the West from CA north to AK and across northern N Amelanchier alnifolia NNAW America

Aruncus dioicus (Aruncus Wet areas of the coastal West from CA Rosaceae sylvester) CWcb north to AK; circumboreal

Comarum palustre Wet areas across northern N America; Rosaceae (Potentilla palustris) NNAcb circumboreal

Crataegus douglasii var. Wet areas of the West from CA north to Rosaceae douglasii NNAW AK and across northern N America

Huggins 65

Introduced throughout N America from Rosaceae Filipendula ulmaria I Eurasia

Wet sand dunes and rocky bluffs of the coastal West from CA north to AK; also S Rosaceae Fragaria chiloensis CWicd America

Wet sand dunes and rocky bluffs of the Fragaria chiloensis ssp. coastal West from CA north to AK; also S Rosaceae lucida CWicd" America

Wet sand dunes and rocky bluffs of the Fragaria chiloensis ssp. coastal West from CA north to AK; also S Rosaceae pacifica CWicd" America

Fragaria virginiana var. Rosaceae glauca NA Widespread throughout N America

Rosaceae Geum calthifolium CBCa Coastal BC north to AK; amphiberingian

Widespread throughout N America; Rosaceae Geum macrophyllum NAa amphiberingian

Geum macrophyllum var. Widespread throughout N America; Rosaceae macrophyllum NAa" amphiberingian

Subalpine and alpine zones of the West Rosaceae Luetkea pectinata W from CA north to AK

Wet lowland areas of the coastal West Rosaceae Malus fusca CW from CA north to AK

Potentilla anserina ssp. Wet areas across northern N America; Rosaceae pacifica NNAcb circumboreal

Beaches and wet areas of the coastal Rosaceae Potentilla egedii CWic West from CA north to AK

Rosaceae Potentilla gracilis NA Widespread throughout N America

Coastal PNW from WA north to AK; Rosaceae Potentilla villosa CPNWa amphiberingian

Rosa nutkana var. Wet areas of the West from CA north to Rosaceae nutkana W AK

Huggins 66

Wet areas across northern N America; Rosaceae Rubus chamaemorus NNAcb circumboreal

Introduced throughout N America from Rosaceae Rubus laciniatus I Eurasia

Widespread throughout the West from CA north to AK; also across northern N Rosaceae Rubus parviflorus NNAW America

Widespread throughout the West from CA Rubus parviflorus north to AK; also across northern N Rosaceae grandiflorus NNAW" America

Widespread throughout the West from CA Rubus parviflorus ssp. north to AK; also across northern N Rosaceae parviflorus NNAW" America

Widespread throughout the PNW from OR Rosaceae Rubus pedatus PNW north to AK

Widespread throughout wet areas of the Rosaceae Rubus spectabilis CPNW coastal PNW from N CA north to AK

Sanguisorba canadensis Wet areas of the montane and subalpine ssp. latifolia (Sanguisorba zones throughout the PNW and E N Rosaceae stipulata) PNWd America

Wet areas of the coastal PNW from WA Rosaceae Sanguisorba menziesii CPNW north to AK

Wet areas of the coastal West from CA Rosaceae Sanguisorba officinalis CW north to AK

Montane, subalpine, and alpine zones of the West from CA north to AK and across Rosaceae Sibbaldia procumbens NNAWcb northern N America; circumboreal

Montane, subalpine, and alpine zones of Rosaceae Sorbus sitchensis PNW the PNW from N CA north to AK

Sorbus sitchensis var. Montane, subalpine, and alpine zones of Rosaceae grayi PNW" the PNW from N CA north to AK

Huggins 67

Widespread in wet areas of the West from Rosaceae Spiraea douglasii W CA north to SE AK

Spiraea douglasii var. Widespread in wet areas of the West from Rosaceae menziesii W" CA north to SE AK

Montane zones of the interior PNW; bad Rosaceae Spiraea pyramidata PNW coordinates?

Wet areas across northern N America; Rubiaceae Galium kamtschaticum NNAa amphiberingian

Rubiaceae Galium trifidum NAcb Wet areas of the northern hemisphere

Galium trifidum ssp. Wet areas of the West from CA north to Rubiaceae columbianum W NW BC

Widespread throughout N America; Rubiaceae Galium triflorum NAcb circumboreal

Ruppia cirrhosa (Ruppia Wet areas throughout N America; Ruppiaceae spiralis) NAcbd circumboreal; also S America

Wet areas throughout N America; Ruppiaceae Ruppia maritima NAcbd circumboreal; also S America

Introduced throughout N America from Salicaceae Salix fragilis I Eurasia

Salix lucida ssp. lasiandra (Salix Widespread in wet areas of the West from Salicaceae lasiandra) W CA north to AK

Widespread in wet areas of the West from Salicaceae Salix scouleriana W CA north to AK

Widespread in wet areas of the West from Salicaceae Salix sitchensis W CA north to AK

Santalaceae Geocaulon lividum NNA Wet areas across northern N America

Acer glabrum var. Widespread throughout the PNW from OR Sapindaceae douglasii PNW north to AK

Widespread throughout the coastal West Saxifragaceae Boykinia occidentalis CW from CA north to NW BC

Huggins 68

Chrysosplenium Wet montane and subalpine zones across Saxifragaceae tetrandrum NNAcb northern N America; circumboreal

Hemieva ranunculifolia (Suksdorfia Montane and subalpine zones of the PNW Saxifragaceae ranunculifolia) PNW from N CA north to NW BC

Wet areas of the montane, subalpine, and alpine zones of the coastal PNW from OR Saxifragaceae Heuchera glabra CPNW north to AK

Widespread throughout the coastal West Saxifragaceae Heuchera micrantha CW from CA north to NW BC

Wet areas of the montane, subalpine, and alpine zones of the coastal PNW from OR Saxifragaceae Leptarrhena pyrolifolia CPNW north to AK

Micranthes ferruginea Wet areas of the West from CA north to Saxifragaceae (Saxifraga ferruginea) W AK

Wet areas of the montane, subalpine, and Micranthes nelsoniana alpine zones of the coastal PNW from OR Saxifragaceae (Saxifraga nelsoniana) CPNWa north to AK; amphiberingian

Wet areas of the montane, subalpine, and Micranthes nelsoniana alpine zones of the coastal PNW from OR Saxifragaceae var. carlottae CPNWa" north to AK; amphiberingian

Wet montane and subalpine zones of the Saxifragaceae Mitella breweri W West from CA north to BC

Saxifraga bronchialis ssp. Subalpine areas from BC north to AK; Saxifragaceae austromontana CBC cordilleran

Widespread throughout the West from CA north to AK and across northern N Saxifragaceae Saxifraga caespitosa NNAWcb America; circumboreal

Wet areas of the West from CA north to Saxifragaceae Saxifraga mertensiana W AK

Huggins 69

Wet areas of the subalpine and alpine zones of the coastal West from CA north Saxifragaceae Saxifraga tolmiei CW to AK

Montane and subalpine zones across Saxifragaceae Saxifraga tricuspidata NNA northern N America

Wet areas of the coastal West from CA Saxifragaceae Tellima grandiflora CW north to AK

Widespread in moist areas of the West Saxifragaceae Tiarella trifoliata W from CA north to AK

Tiarella trifoliata var. laciniata (Tiarella Saxifragaceae laciniata) CPNW" Coastal PNW from WA OR north to AK

Widespread in wet areas of the West from Tiarella trifoliata var. CA north to AK; more common at lower Saxifragaceae trifoliata W" elevations

Widespread in wet areas of the West from Tiarella trifoliata var. CA north to AK; more common at higher Saxifragaceae unifoliata W" elevations

Widespread in wet areas of the coastal Saxifragaceae Tolmiea menziesii CW West from CA north to AK

Infrequent in wet areas of the West from CA north to AK and across northern N Scrophulariaceae Limosella aquatica NNAWcb America; circumboreal

Wet areas across northern N America; Selaginellaceae Selaginella selaginoides NNAcb circumboreal

Wet areas of the PNW from N CA north to Selaginellaceae Selaginella wallacei PNW NW BC

Taxaceae Taxus brevifolia CPNW Coastal PNW from N CA north to SE AK

Phegopteris connectilis Wet areas throughout N America; also Thelypteridaceae (Thelypteris phegopteris) NAd Eurasia

Huggins 70

Thelypteris quelpaertensis (Thelypteris Wet areas of the coastal PNW from WA limbosperma) north to AK; amphiberingian; disjunct in Thelypteridaceae (Thelypteris oreopteris) CPNWad NE N America

Triantha glutinosa Tofieldiaceae (Tofieldia glutinosa) NNA Wet areas across northern N America

Triantha occidentalis ssp. Wet areas of the PNW from OR north to Tofieldiaceae brevistyla PNW AK

Wet areas of the West from CA north to Sparganium AK and across northern N America; Typhaceae angustifolium NNAWcb circumboreal

Wet areas of the West from CA north to AK and across northern N America; Typhaceae Sparganium emersum NNAWcb circumboreal

Wet areas of the West from CA north to Sparganium emersum AK and across northern N America; Typhaceae ssp. emersum NNAWcb" circumboreal

Sparganium Wet areas across northern N America; Typhaceae hyperboreum NNAcb circumboreal

Wet areas of the West from CA north to Sparganium natans AK and across northern N America; Typhaceae (Sparganium minimum) NNAWcb circumboreal

Widespread in wet areas of the West from Violaceae Viola glabella W CA north to AK

Wet areas of the coastal PNW from OR Violaceae Viola langsdorfii CPNWa north to AK; amphiberingian

Intertidal zones of the West from MX north Zosteraceae Phyllospadix scouleri Wic to AK

Intertidal zones of the PNW from OR north Zosteraceae Phyllospadix serrulatus PNWic to AK

Huggins 71

Intertidal zones of the West from MX north Zosteraceae Phyllospadix torreyi Wic to AK

Zostera marina (Zostera Zosteraceae angustifolia) Cic Cosmopolitan species of intertidal zones

Huggins 72

Appendix C: Vascular plant inventory of Pitt Island, BC

Table 5: Informal inventory of vascular plant species in study area on Pitt Island, BC.

Family Genus Species Common Name

Adoxaceae Sambucus racemosa Red Elderberry

Apiaceae Angelica lucida Sea­watch

Apiaceae Conioselinum pacifica Pacific Hemlock­Parsley

Apiaceae Ligusticum calderi Calder's Mountain Lovage

Apiaceae Ligusticum scoticum Beach Lovage

Apiaceae Osmorhiza purpurea Purple Sweet­Cicely

Araceae Lysichiton americanus Skunk Cabbage

Araliaceae Oplopanax horridus Devil's Club

Asparagaceae Maianthemum dilatatum False Lily­of­the­Valley

Asteraceae Achillea millefolium Yarrow

Asteraceae Arnica latifolia Broadleaf Arnica

Asteraceae Artemisia norvegica ssp. saxatilis Mountain Sagewort

Asteraceae Hieracium triste Hairy Hawkweed

Asteraceae Microseris borealis Apargidium

Asteraceae Nabalus alatus Western Rattlesnake Root

Asteraceae Petasites frigidus ssp. nivalis Sweet Coltsfoot

Asteraceae Senecio triangularis Arrowleaf Groundsel

Athyriaceae Athyrium filix­femina Lady Fern

Betulaceae Alnus rubra Red Alder

Betulaceae Alnus viridis ssp. sinuata Sitka Alder

Blechnaceae Blechnum spicant Deer Fern

Brassicaceae Cardamine oligosperma var. kamtschatica Little­seeded Bittercress

Brassicaceae Cochlearia groenlandica Scurvy Grass

Huggins 73

Caprifoliaceae Valeriana sitchensis Sitka Valerian

Caryophyllaceae Sagina maxima Coastal Pearlwort

Caryophyllaceae Spergularia canadensis Canadian Sand­spurry

Caryophyllaceae Stellaria crispa Crisp Starwort

Caryophyllaceae Stellaria humifusa Salt Marsh Starwort

Cornaceae Cornus canadensis Bunchberry

Cupressaceae Chamaecyparis nootkatensis Yellow Cedar

Cupressaceae Juniperus communis Common Juniper

Cupressaceae Thuja plicata Western Redcedar

Cyperaceae Carex circinata Coiled Sedge

Cyperaceae Carex macrochaeta Big­awned Sedge

Cyperaceae Carex mertensii Merten's' Sedge

Cyperaceae Carex lyngbyei Salt Marsh Sedge

Cyperaceae Eriophorum angustifolium Narrow Cotton Grass

Cyperaceae Trichophorum caespitosum Tufted Clubrush

Cystopteridaceae Cystopteris fragilis Fragile Fern

Cystopteridaceae Gymnocarpium dryopteris Oak Fern

Dennstaedtiacea e Pteridium aquilinum Bracken Fern

Droseraceae Drosera rotundifolia Round­leaved Sundew

Dryopteridaceae Dryopteris expansa Shield Fern

Dryopteridaceae Dryopteris filix­mas Male Fern

Dryopteridaceae Polystichum munitum Sword Fern

Ericaceae Andromeda polifolia Bog Rosemary

Ericaceae Cassiope mertensiana White Mountain­Heather

Ericaceae Elliottia pyrolifolia Copperbush

Ericaceae Empetrum nigrum Crowberry

Ericaceae Gaultheria shallon Salal

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Ericaceae Harrimanella stelleriana Alaska Mountain­Heather

Ericaceae Kalmia microphylla Western Bog­Laurel

Ericaceae Kalmia procumbens Alpine­Azalea

Ericaceae Menziesia ferruginea False Azalea

Ericaceae Moneses uniflora Wax­

Ericaceae Phyllodoce glanduliflora Yellow Mountain­Heather

Ericaceae Rhododendron groenlandicum Labrador Tea

Ericaceae Vaccinium alaskense Alaska Blueberry

Ericaceae Vaccinium caespitosum Dwarf Blueberry

Ericaceae Vaccinium ovalifolium Oval­leaf Blueberry

Ericaceae Vaccinium oxycoccus Bog Cranberry

Ericaceae Vaccinium parvifolium Red Huckleberry

Ericaceae Vaccinium uliginosum Bog Blueberry

Ericaceae Vaccinium vitis­idaea Lingonberry

Fabaceae Lupinus nootkatensis Nootka Lupine

Gentianaceae Gentiana douglasiana Swamp Gentian

Gentianaceae Gentiana platypetala Broad­leaved Gentian

Grossulariaceae Ribes bracteosum Stink Currant

Grossulariaceae Ribes laxiflorum Trailing Currant

Lentibulariaceae Pinguicula vulgaris Common Butterwort

Liliaceae Fritillaria camschatcensis Rice Root

Liliaceae Streptopus amplexifolius Clasping Twisted­Stalk

Liliaceae Streptopus lanceolatus var. roseus Rosy Twisted­Stalk

Lycopodiaceae Diphasiastrum alpinum Alpine Clubmoss

Lycopodiaceae Diphasiastrum complanatum Ground­Cedar

Lycopodiaceae Huperzia haleakale (selago) Fir Clubmoss

Lycopodiaceae Lycopodium annotinum Stiff Clubmoss

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Lycopodiaceae Lycopodium clavatum Running Clubmoss

Lycopodiaceae Lycopodium dendroideum Ground­Pine

Lycopodiaceae Lycopodium sitchense Alaska Clubmoss

Melanthiaceae Veratrum viride Corn Lily

Nephrophyllidiu Menyanthaceae m crista­galli Deer Cabbage

Myricaceae Myrica gale Sweet Gale

Onagraceae Epilobium anagallidifolium Small Willowherb

Onagraceae Epilobium angustifolium Fireweed

Orchidaceae Listera caurina Broad­leaf Twayblade

Orchidaceae Listera cordata Heartleaf Twayblade

Northern Green Bog Orchidaceae Platanthera aquilonis Orchid

Orobanchaceae Castilleja miniata Common Red Paintbrush

Orobanchaceae Castilleja parviflora ssp. oreopola Magenta Paintbrush

Orobanchaceae Pedicularis ornithorhyncha Birds­beak Lousewort

Pinaceae Abies amabilis Amabilis Fir

Pinaceae Picea sitchensis Sitka Spruce

Shore Pine / Lodgepole Pinaceae Pinus contorta Pine

Pinaceae Tsuga heterophylla Western Hemlock

Pinaceae Tsuga mertensiana Mountain Hemlock

Plantaginaceae Penstemon davidsonii Davidson's Penstemon

Plantaginaceae Plantago maritima Sea Plantain

Poaceae Calamagrostis canadensis Bluejoint Reedgrass

Poaceae Deschampsia elongata Slender Hairgrass

Polypodiaceae Polypodium glycyrrhiza Licorice Fern

Portulacaceae Claytonia sibirica Siberian Miner's Lettuce

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Tall Mountain Primulaceae Dodecatheon jeffreyi Shooting­Star

Primulaceae Trientalis europaea ssp. arctica Arctic Starflower

Ranunculaceae Anemone narcissiflora Narcissus Anemone

Ranunculaceae Aquilegia formosa Western Columbine

Ranunculaceae Caltha leptosepala Marsh­Marigold

Ranunculaceae Coptis asplenifolia Few­leaved Goldthread

Ranunculaceae Coptis trifolia Three­leaf Goldthread

Ranunculaceae Kumlienia cooleyae Cooley's False Buttercup

Ranunculaceae Ranunculus uncinatus Small­flowered Buttercup

Rosaceae Aruncus dioicus Goatsbeard

Rosaceae Geum calthifolium Caltha­leaved Avens

Rosaceae Luetkea pectinata Partridge­Foot

Rosaceae Malus fusca Pacific Crabapple

Rosaceae Potentilla anserina ssp. pacifica Silverweed

Rosaceae Rosa nutkana Nootka Rose

Rosaceae Rubus parviflora Thimbleberry

Rosaceae Rubus pedatus Five­leaf Bramble

Rosaceae Rubus spectabilis Salmonberry

Rosaceae Sanguisorba canadensis Sitka Burnett

Rosaceae Sorbus sitchensis Sitka Mountain Ash

Rubiaceae Galium trifidum Small Bedstraw

Saxifragaceae Heuchera glabra Smooth Alumroot

Saxifragaceae Leptarrhena pyrolifolia Leatherleaf Saxifrage

Saxifragaceae Micranthes nelsoniana Heartleaf Saxifrage

Saxifragaceae Saxifraga ferruginea Rusty­leaf Saxifrage

Saxifragaceae Saxifraga tolmiei Tolmie's Saxifrage

Saxifragaceae Tellima grandifolia Fringecups

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Saxifragaceae Tiarella trifoliata var. trifoliata Foamflower

Saxifragaceae Tiarella trifoliata var. unifoliata One­leaf Foamflower

Taxaceae Taxus brevifolia Pacific Yew

Thelypteridaceae Phegopteris connectilis Narrow Beech Fern

Tofieldiaceae Triantha glutinosa Sticky False­Asphodel

Violaceae Viola glabella Stream Violet

Violaceae Viola langsdorfii Alaska Violet

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Appendix D: Generalized Extent of Floristic Elements

Figure 10: Generalized representation of floristic elements employed in the local and regional inventories.

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Appendix E: Vascular Plant Inventory of Hevenor Islet, BC

Table 6: Informal inventory of vascular plant species on Hevenor Islet, BC; graminoids not included.

Family Genus Species Common Name Adoxaceae Sambucus racemosa Red Elderberry Asparagaceae Maianthemum dilatatum False Lily-of-the-Valley Athyriaceae Athyrium filix-femina Lady Fern Cupressaceae Thuja plicata Western Redcedar Dryopteridaceae Dryopteris expansa Shield Fern Ericaceae Gaultheria shallon Salal Ericaceae Vaccinium alaskense Alaska Blueberry Ericaceae Vaccinium ovalifolium Oval Leaf Blueberry Ericaceae Vaccinium parvifolium Red Huckleberry Grossulariaceae Ribes laxiflorum Trailing Currant Pinaceae Picea sitchensis Sitka Spruce Polypodiaceae Polypodium glycyrrhiza Licorice Fern Ranunculaceae Ranunculus uncinatus Small-flowered Buttercup Rosaceae Malus fusca Pacific Crabapple Rosaceae Rubus parviflorus Thimbleberry Rosaceae Rubus spectabilis Salmonberry Saxifragaceae Micranthes nelsoniana Nelson's Saxifrage Saxifragaceae Tellima grandiflora Fringecups Taxaceae Taxus brevifolia Pacific Yew