Plant Species Coalition Groups of Zion National Park: an Individualistic, Floristic Alternative to Vegetation Classification Jeffrey E

Monographs of the Western North American Naturalist

Volume 8 Article 2

5-28-2015 Plant species coalition groups of Zion National Park: an individualistic, floristic alternative to vegetation classification Jeffrey E. Ott USDA Forest Service, Rocky Mountain Research Station, Boise, ID, [email protected]

Stewart C. Sanderson USDA Forest Service, Rocky Mountain Research Station, Provo, UT, [email protected]

E. Durant McArthur USDA Forest Service, Rocky Mountain Research Station, Provo, UT, [email protected]

Follow this and additional works at: https://scholarsarchive.byu.edu/mwnan

Recommended Citation Ott, Jeffrey E.; Sanderson, Stewart C.; and McArthur, E. Durant (2015) "Plant species coalition groups of Zion National Park: an individualistic, floristic alternative to vegetation classification," Monographs of the Western North American Naturalist: Vol. 8 , Article 2. Available at: https://scholarsarchive.byu.edu/mwnan/vol8/iss1/2

This Monograph is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Monographs of the Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Monographs of the Western North American Naturalist 8, © 2015, pp. 26–97

PLANT SPECIES COALITION GROUPS OF ZION NATIONAL PARK: AN INDIVIDUALISTIC, FLORISTIC ALTERNATIVE TO VEGETATION CLASSIFICATION

Jeffrey E. Ott1, Stewart C. Sanderson2 and E. Durant McArthur2

ABSTRACT.—Vegetation surveys at Zion National Park (Zion), Utah, have contributed to our understanding of plant community patterns and their relationship to environmental factors. Previous authors used vegetation plot data to char- acterize vegetation types at Zion following conventional procedures that emphasize spatial discreteness and dominant species. We developed and applied an alternative approach for community characterization that emphasizes nondiscrete presence-absence patterns and is compatible with the individualistic concept. We reanalyzed existing plot data from Zion using coalition clustering, an algorithm that identifies groups of positively-associated species referred to as coali- tion groups. Each species and plot in the data set was linked to each coalition group via an “affinity” value obtained through weighted averaging. Affinity values were used to characterize environmental affinities of coalition groups through regression tree modeling and predictive mapping. We also identified species that frequently co-occurred with coalition groups (affiliate species) and those that frequently co-occurred with high cover (dominant-affiliates), viewing these as alternatives to conventional prevalent and dominant species. Following this approach, we identified 10 coalition groups at Zion that overlapped compositionally and spatially to differing degrees. Mesic environments on a gradient from low-elevation riparian zones through mid-elevation narrow canyons to high-elevation plateaus were represented by 3 overlapping groups. Two groups occupying slickrock and sand environments were detected on the Navajo Sand- stone, as well as 2 on mesa tops above it. At lower elevations, 3 intergrading xeric coalition groups were distinguished. When previously classified associations of the National Vegetation Classification were clustered based on shared affini- ties to coalition groups, the arrangement differed from existing classification schemes but was environmentally inter- pretable. Although these patterns are contingent on conditions at the time of data collection, they provide a baseline that could be used for evaluating and predicting plant community change in the park. With proper attention to sampling and analysis issues, our community characterization approach could be applied in other settings as an alternative or sup- plement to conventional vegetation classification.

RESUMEN.—Monitoreos de la vegetación en el Parque Nacional Zion (Zion), Utah, han contribuido a nuestro entendimiento de los patrones de las comunidades de plantas y su relación con factores ambientales. En estudios anteri- ores, los autores utilizaron datos de terrenos con vegetación para caracterizar tipos de vegetación en Zion, siguiendo procedimientos convencionales que enfatizan estados discretos espaciales y especies dominantes. Desarrollamos y apli- camos un enfoque alternativo para la caracterización de la comunidad que enfatiza los patrones de presencia-ausencia no-discreta y es compatible con el concepto individualista. Reanalizamos datos gráficos de vegetación de Zion utilizando agrupamiento de coalición, un algoritmo que identifica grupos de especies positivamente asociadas, llamados grupos de coalición. Cada especie y terreno en el conjunto de datos estaba unido a cada grupo de coalición por medio de un valor de “afinidad” obtenido a través de un promedio ponderado. Los valores de afinidad se emplearon para caracterizar afinidades ambientales de grupos de coalición a través de árboles de regresión y mapeo predictivo. También identifi- camos especies que frecuentemente coexistían con grupos de coalición (especies afiliadas) y aquellas que frecuente- mente coexistían con cubiertas altas (afiliados dominantes), considerándolas como alternativas a las especies prevalentes y dominantes. Siguiendo este enfoque, identificamos diez grupos de coalición en Zion que se superponían en composi- ción y espacialmente en diversos grados. Ambientes relativamente húmedos en un gradiente de baja elevación, zona riparia, a elevación media, cañones estrechos, hasta elevación alta, meseta, estuvieron representadas por tres grupos sobrelapados. Dos grupos que ocupaban medios arenosos y con dunas fueron detectados en Navajo Sandstone, así como dos en la cima de la meseta por encima de esa zona. En elevaciones más bajas, se distinguieron tres grupos de coalición de interacción árida. Cuando se agruparon las asociaciones previamente clasificadas de la Clasificación Vegetal Nacional basándose en las afinidades compartidas con los grupos de coalición, la disposición se diferenció de los esquemas de clasificación existentes, pero eran interpretables según el ambiente. A pesar de que estos patrones dependen de las condiciones al momento de la recolección de datos, proporcionan un punto de referencia que se podría utilizar para evaluar y predecir cambios en la comunidad vegetal en el parque. Con la atención adecuada a las cuestiones del muestreo y análisis, nuestro enfoque de caracterización de la comunidad se podría aplicar en otros ambientes como una alternativa o suplemento a la clasificación de vegetación convencional.

1USDA Forest Service, Rocky Mountain Research Station, Boise, ID 83702. E-mail: [email protected] 2USDA Forest Service, Rocky Mountain Research Station, Provo, UT 84606.

26 2015] PLANT COALITION GROUPS OF ZION NP 27

Zion National Park (Zion), Utah, is best Harper et al. (2001) and Cogan et al. (2004) known for its striking geological features but followed conventional vegetation classification also boasts a high diversity of plant taxa and practices of delineating spatially discrete vege - plant communities. Over 800 native vascular tation types differentiated primarily by domi- plant species (Fertig and Alexander 2009, Fer- nant species or vegetation structure. While tig et al. 2012) and 95 vegetation associations these practices are widely followed and useful (Cogan et al. 2004) are distributed across for many purposes (Mueller-Dombois and pronounced edaphic and climatic gradients Ellenberg 1974, Grossman et al. 1998), they within the park (Harper 1993, Biek et al. 2003, result in community characterizations that Cogan et al. 2004, O’Meara 2006, Fowler et emphasize certain ecological patterns at the al. 2007, Fertig and Alexander 2009). The con- possible expense of others. Continuous grada- centrated and contrasting environmental and tions of community composition due to indi- botanical diversity of Zion contribute to the vidualistic species responses (Gleason 1926, park’s scientific and conservation value. Given 1939) can be obscured by discrete community the U.S. National Park Service mandate to models, and distributions of subordinate species protect natural resources for the enjoyment of might not be well represented by those of present and future generations, information dominants. These issues are relevant at Zion on the distribution, composition and environ- because species responses to the park’s com- mental attributes of plant communities at plex overlay of environmental gradients are Zion is important for science-based resource likely to be individualistic, and because the management (NPS 2006). number of structurally or numerically domi- Numerous workers have contributed to nant species is small relative to the number of current knowledge of vascular plant communi- species in the flora as a whole (Cogan et al. ties at Zion. Plant collection efforts from 1894 2004, Fertig and Alexander 2009). to the present have documented the flora of In this paper we present an alternative the park and provided insights concerning plant community characterization for Zion that species distributions and habitat relations (re- mirrors the earlier vegetation classification of viewed in Fertig and Alexander 2009, Fertig Harper (1993) but is neither spatially discrete et al. 2012). Woodbury’s (1933) monograph of nor built around dominant species. Instead biotic communities was followed by more of following Harper’s (1993) approach using detailed descriptions of specific plant com- dominant species as classification criteria, then munities such as hanging gardens (Malanson characterizing the floristic composition of 1980, Malanson and Kay 1980, Welsh 1989, communities thus classified, we essentially re - Fowler et al. 2007) and isolated mesa tops versed the process by first identifying species (Madany and West 1983, 1984). Two park-wide, groups defined by presence-absence patterns, plot-based vegetation surveys have been car- then identifying dominant species and other- ried out at Zion, one in 1987–1989 (Harper wise characterizing the communities and envi- 1993, Harper et al. 2001) and the other in ronments where these species groups occur. 1999–2003 (Cogan et al. 2004). Harper (1993) This was accomplished by reanalyzing the and Harper et al. (2001) identified 10 major Harper (1993) data set using coalition cluster- vegetation types at Zion using an informal, ing and other algorithms from the RCLUS expert-knowledge approach, while Cogan et computer program (Sanderson et al. 2006) in al. (2004) characterized vegetation more for- combination with regression tree modeling mally and in greater detail according to the (Breiman 1993) and predictive mapping (Fer- U.S. National Vegetation Classification (NVC) rier and Guisan 2006). Because RCLUS is (Grossman et al. 1998, FGDC 2008, Jennings relatively new and its algorithms are here ap - et al. 2009). These studies have jointly yielded plied in a novel way, one of our objectives is an enviable amount of information but they to outline the conceptual and methodological have not exhausted all that can be known basis of our approach. We draw conceptual about plant communities at Zion. In this paper parallels that align our approach with Curtis’s we demonstrate that even in the absence of (1959) modal and prevalent species concepts, new data, alternative approaches for data ana ly - applied here in a spatially nondiscrete way. sis and synthesis can provide valuable new We demonstrate the ecological insights gained information and insights. from our approach by comparing our results 28 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 directly with those of Harper et al. (2001) and entisols, aridisols, mollisols, and alfisols (Morten - Cogan et al. (2004). We also use our results to sen 1977). Some soils have distinctive physical build an alternative higher-level classification and chemical properties, as exemplified by the of the NVC associations described by Cogan bentonite shrink-swell clays of the Petrified et al. (2004) that reflects their position in rela- Forest member of the Chinle Formation (Biek tion to environmental gradients at Zion. et al. 2003). Cryptobiotic crusts are present on many soil surfaces and contribute to soil sta- METHODS bility and fertility (cf. Belnap et al. 2000). Although most of Zion currently has an Study Area undeveloped wilderness character, vegetation Zion National Park (Zion), Utah, USA, covers has been affected by historic floodplain agri- approximately 590 km2 at the western edge of culture, livestock grazing, and fire suppression, the Colorado Plateau physiographic/floristic resulting in increases in invasive annuals and area, bordering the Great Basin to the north- woody plants at the expense of native grasses west and the Mojave Desert to the southwest and forbs (Woodbury 1933, Madany and West (Biek et al. 2003, McLaughlin 1986). Climate 1983, Cogan et al. 2004). Although livestock at Zion is mostly semiarid and temperate but numbers have dropped since the early twenti- varies greatly across an elevation gradient of eth century, mule deer (Odocoileus hemionus) 1128–2660 m (Cogan et al. 2004). Precipita- have increased in areas of high human visita- tion is concentrated in winter/spring frontal tion, resulting in declines in cottonwood (Popu- storms (bringing snow to higher elevations) lus fremontii) and other riparian species and late-summer monsoons (Woodbury 1933, (Ripple and Beschta 2006). The troublesome Mortensen 1977). At Zion Canyon (1234 m), invasive species cheatgrass (Bromus tectorum), weather records from 1904 to 2012 show a red brome (Bromus rubens), ripgut brome (Bro- mean annual precipitation of 382 mm and mean mus diandrus), and tamarisk (Tamarix spp.) were temperatures of 5 °C in January and 29 °C in already well-established in Zion at the time July (WRCC 2014). of the Harper (1993) and Cogan et al. (2004) Horizontally aligned exposures of sedimen- vegetation surveys. tary rocks are prominent at Zion. The oldest Vegetation Data strata occur at low elevations on the south- western side of the park and on slightly higher Our primary data source is the vegetation elevations on the northwestern side where survey carried out in 1987–1989 by Dr. K.T. the Taylor Creek thrust fault disrupts their Harper and associates (Harper 1993, Harper stratigraphic continuity (Biek et al. 2003, et al. 1992, 2001, 2003), which we refer to O’Meara 2006). The lowermost geologic for- as the Harper Survey. This was the first mations (Kaibab, Moenkopi, Chinle, Moen- park-wide, plot-based vegetation survey con- have, and Kayenta) contain alternating bands ducted for Zion. The Harper survey followed a of mudstone, siltstone, sandstone, conglomer- systematic sampling strategy in which U.S. ate, limestone, and gypsum that erode into Geological Survey (USGS) cadastral section badlands, slopes, and cliffs (Biek et al. 2003). corners and corresponding park boundary These strata are dwarfed by the larger cliffs markers served as sampling targets (Fig. 1A). and ‘slickrock’ exposures of the overlying In addition to 270 plots located at or near Navajo Sandstone (the park’s primary scenic these markers, 18 plots were placed using attraction), which is further overlain by sand- subjective criteria so as to capture spatially stone, limestone, shale, and gypsum deposits restricted plant communities (riparian areas, of the Temple Cap, Carmel, and Cedar Moun- hanging gardens, and old fields). Data col- tain formations (Biek et al. 2003, O’Meara lected within 100-m2 circular plots (or rectan- 2006). The sedimentary sequence is in turn gular, in 6 cases) included vascular plant species partially covered by alluvial, lacustrine, land- composition following Welsh et al. (1987), ocu- slide, talus, residual, eolian, and volcanic (basalt lar estimates of species cover classes according flows and cinder cones) deposits (Biek et al. to the Braun-Blanquet scale (Braun-Blanquet 2003, O’Meara 2006). 1932), and percent cover of bare soil, rock, Soils at Zion are generally poorly developed litter, and cryptogams (Harper 1993, Harper and have been classified as rock outcrops, et al. 2001). 2015] PLANT COALITION GROUPS OF ZION NP 29

Fig. 1. Plot locations in Zion National Park, Utah, as sampled by 2 vegetation surveys referenced in this paper.

Harper (1993) used the Harper survey data hereafter USGS Survey, which featured 346 to classify the vegetation of Zion into ten vege- circular, square, or rectangular plots (Fig. 1B) tation types that were concisely summarized ranging from 100 m2 to 400 m2 where plant in Harper et al. (2001). Plots were assigned species were identified following Kartesz to vegetation types “commonly used by land (1999). A gradsect strategy (Gillison and managers in the area” based on dominant Brewer 1985) was used to divide Zion into species and environmental context (Harper biophysical units (BPUs) that were mapped et al. 2001). Prevalent and modal species and targeted for field sampling. Accessible were characterized for each vegetation type polygons of each BPU were targeted by field following Curtis (1959). Prevalent species are survey teams who placed plots on sites defined as the most frequently encountered deemed representative of major vegetation species (based on plot occurrences) within a types and relatively homogeneous in terms community or vegetation type, where the num - of vegetation structure and composition ber of prevalent species is equal to the species (Cogan et al. 2004). Plot data were collected density (average number of species per plot) in 1999–2000. for the community (Curtis 1959, Harper et Cogan et al. (2004) used the USGS survey al. 2001). Modal species are defined as preva- data to classify and map the vegetation of Zion lent species that reach their greatest fre- following the U.S. National Vegetation Classi- quency in a given community relative to other fication System (NVC) (FGDC 2008, Jennings communities under study. In Harper et al. et al. 2009). Each plot was assigned to an asso- (2001), modal species were identified for each ciation nested within higher levels of the NVC vegetation type relative to the others de - as defined at the time of publication (Gross- scribed at Zion. man et al. 1998, Cogan et al. 2004). Multivari- The Hanging Gardens vegetation type ate ana lysis of species cover class data by described by Harper (1993) and Harper et strata guided the classification process (Cogan al. (2001) incorporated data from published et al. 2004). studies of Malanson and Kay (1980) and Welsh We merged the Harper data set, the list (1989); however, these data were excluded of prevalent and modal species by vegeta- from the current analysis, which uses only plot tion type from Harper et al. (2001), the USGS data from the actual Harper survey. data sets containing species composition and We also draw from plot data originating NVC classification by plot, and our RCLUS with the USGS–NPS Vegetation Characteri - results (described below) into a single data- zation Program at Zion (Cogan et al. 2004), base. Taxonomic names from these sources 30 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 were standardized to Welsh et al. (2003) total number of occurrences in the table (Jack- nomenclature so that they could be matched son et al. 1989, Tichý and Chytrý 2006). from one data set to another. Coalition clustering agglomerates species whose mean pairwise association is higher Coalition Clustering than the user-defined threshold. The algorithm We used coalition clustering, a nonhierar- initiates the first group with a randomly se- chical agglomerative clustering algorithm in- lected species and then randomly cycles through cluded in the program RCLUS (Sanderson et al. the remaining species, adding them to existing 2006), to identify groups of species with simi- groups or initiating new groups as appropriate. lar patterns of presence-absence in the Harper Species can belong to multiple groups with data set. RCLUS was developed as a tool for the exception that species already present in R-mode (species-based) clustering of commu- an existing group cannot initiate a new group. nity data sets. The name coalition clustering The algorithm undergoes several cycles of ad - underscores the individualistic variability of ding species, removing species and merging species within the clusters (coalitions) that the clusters allowing the composition of species algorithm builds. Coalition clustering gener- groups to stabilize. The threshold association ates clusters of positively associated species value affects group number and size; high here referred to as coalition groups. The term thresholds tend to produce many small groups core species, used by Sanderson et al. (2006) to in contrast to few large groups at low thresh- refer to species belonging to a group, is here olds (Sanderson et al. 2006). replaced by the term coalition species. Once the Coalition Clustering algorithm is Different coalition clustering solutions can complete, RCLUS saves the mean associa- be obtained by varying 3 user-defined parame - tion—here referred to as affinity—of each ters of the algorithm: association coefficient, species to each group (Sanderson et al. 2006). threshold association value, and minimum num - Affinity values provide a quantitative mea - ber of occurrences for inclusion in groups. sure of the relationship between individual Clustering solutions may also vary from one species and coalition groups (calculated for all run of the algorithm to another because of the species, not just those belonging to groups). random nature of the agglomerative process RCLUS also averages affinity values for each (Sanderson et al. 2006). The clustering solu- group across all species in each plot, yielding tion presented here was selected from among values that can be correlated with environ- a small number of variants obtained using the mental variables at plot locations (Sanderson phi coefficient at a threshold value of 0.15, et al. 2006). Our use of the term affinity thus allowing species occurring in 3 or more plots has a specific quantitative meaning that en - to be included in groups. compasses 3 contexts: species affinities to Association coefficients measure the degree species groups (mean pairwise association to which 2 species show the same pattern of with members of the group), spatial affinities presence and absence in a data set, calculated of species groups (plot-averaged species affini- from the cells of a two-way presence-absence ties), and environmental af finities of species contingency table: groups (environmental correlates of spatial affinities). Species 1 Affiliation and Dominance Calculations Present Absent Present a b In addition to generating coalition groups, Species 2 Absent c d RCLUS contains a function that calculates percent co-occurrence of each species to each The phi coefficient incorporates all 4 cells of coalition group. Percent co-occurrence is the table according to the formula calculated by summing joint presences across all pairings of a focal species with the species ______ad – bc comprising a coalition group and then stan- phi coefficient = ______. √(a+b)(a+c)(b+d)(c+d) dardizing by the maximum possible number of joint presences. Using the notation of Phi ranges from –1 (negative association) to +1 presence-absence frequencies, this can be (positive association) and is independent of the represented as follows: 2015] PLANT COALITION GROUPS OF ZION NP 31

Focal species are proxies for the major physical and chemi- Present Absent cal gradients affecting plant distributions at Coalition species Present a b Zion. GIS layers for these variables were available or easily created and integrated in spN a ArcGIS (ESRI 2003, 2004) (Appendixes 1, 2). ∑ Our use of GIS-based variables permitted Percent co-occurrence (affiliation) = ______sp1 spN predictive mapping of species group affinities ∑ (a+b) beyond plot locations (Ferrier and Guisan 2006). sp1 Plot coordinates from the Harper Survey, originally marked on paper maps, were digi- Thus percent co-occurrence of the focal species tized manually with the help of a high-resolu- to the coalition group is the sum of joint pres- tion aerial orthophoto of Zion National Park ences between the focal species and each of (Cogan et al. 2004), digital USGS topographic the coalition species in the group (sp1 to spN), maps, and cadastral survey marker coordinates standardized by the total number of occur- acquired through the USDI Bureau of Land rences of all the coalition species. We have Management (BLM 2006). Hawth’s Tools (Beyer adopted the term affiliation to use inter- 2004) was used to extract environmental data changeably with percent co-occurrence. values for each plot based on its coordinates. Using percent co-occurrence values from Elevation and other topographic variables RCLUS, we identified a set of species with were obtained from a 10-m digital elevation high affiliation (affiliate species) for each model (DEM) acquired from the U.S. National coalition group. Given that Curtis (1959) used Park Service (NPS, personal communication). species density, the mean number of species Slope and exposure were calculated using the per plot in a community, to determine how slope and hillshade functions, respectively, in many species to include in prevalent species ArcGIS Spatial Analyst (ESRI 2004) (Appen- lists, we followed an analogous procedure for dix 1). Hillshade was implemented at 215° determining the number of affiliate species for azimuth and 45° altitude such that exposure each coalition group. The species density of a values were highest on steep southwest-facing coalition group was calculated as the mean slopes where solar heat loading is expected to number of species occurring in plots where be highest (cf. McCune and Keon 2002). Topo- coalition species occurred, weighting plots by graphic position was defined using the how many coalition species they contained. Topographic Position Index (TPI), calculated Extending the affiliate species concept fur- as the difference in elevation between a pixel ther to include abundance as well as occur- and the mean of surrounding pixels within a rence, we calculated the abundance of each defined radius (Weiss 2001). We created 2 TPI affiliate species as its mean cover across plots layers at radii of 50 m (Appendix 1) and 100 m. containing coalition species, weighted once A 1:24,000-scale digital geologic map built again by the number of coalition species per from multiple maps of the Utah Geologic Sur- plot. Arithmetic midpoints of cover classes vey (O’Meara 2006) was converted to raster were used in these calculations. In this way format at the same resolution as the DEM and we were able to identify the dominant species simplified from about 100 map units to 16 occurring alongside members of each coalition geologic substrate classes (Table 1, Appendix 2). group. Affiliate species with mean cover >2% We merged units with similar lithology, spatial were considered dominant-affiliate for our continuity, and/or topographic characteristics, purposes. guided by exploratory analyses of relationships between putative substrate groupings and coali- Environmental Modeling and Mapping tion group affinities. Our groupings include Plot-averaged affinity values of coalition each of the major geologic formations of the species groups, obtained as described above park (Moenkopi [+Kaibab], Chinle, Moenhave, for each plot of the Harper survey, were inte- Kayenta, Navajo, Temple Cap, and Carmel grated with environmental variables for cor- [+Cedar Mountain]) and Quaternary deposits relative modeling. We selected 5 variables— grouped according to their dominant deposi- elevation, slope, topographic exposure, topo- tional material (alluvium, colluvium, eolian, graphic position, and geologic substrate—that lacustrine, mass movement, residuum, and 32 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

TABLE 1. Geologic substrate classes used for environmental modeling of plant coalition group affinities at Zion National Park, Utah. For each substrate class, corresponding map units from a digital geologic map of the Utah Geologi- cal Survey (O’Meara 2006) are shown in the rightmost column. Code Substrate class Descriptiona Map unitsb All Alluvium river/stream deposits Qa*, Qf Bas Basalt volcanics (flows, cinders, ash) Qb*, Qms(b) Car Carmel (+Cedar Mountain) limestone, shale, sandstone, Jc*, Kcmc Formation siltstone, gypsum Chi Chinle Formation mudstone, claystone, siltstone, TRc* sandstone, conglomerate Col Colluvium unconsolidated fine gravity deposits Qc* Eol Eolian unconsolidated wind deposits Qe* Kay Kayenta Formation siltstone, sandstone, mudstone Jk* Lac Lacustrine lake deposits Ql* Mca Mass movement/colluvium/ massive gravity deposits mixed with fine Qmcp* alluvial pediment-mantle gravity and river/stream deposits Mnk Moenkopi (+Kaibab) siltstone, mudstone, limestone, sandstone, TRm*, Pk* Formation conglomerate, gypsum, dolomite Mnv Moenave Formation mudstone, sandstone, siltstone Jm* Nav Navajo Sandstone sandstone Jn* Res Residuum deposits from in situ weathering Qr* Sli Slide/slump/flow massive gravity deposits Qmfy, Qms* (slides, slumps, and flows) Tal Talus coarse gravity deposits Qmt*, QTng Tem Temple Cap Formation sandstone, siltstone, mudstone Jt* aSubstrate description summarized from Biek et al. (2003) and O’Meara (2006) bMap units as coded in O’Meara (2006) *Wildcard indicating multiple map units containing the listed prefix volcanics). Mass movement deposits were fur- For the Harper et al. (2001) data set, we calcu- ther split into 3 classes that occupy different lated mean affinity values across modal species topographic settings in the park: (1) talus, (2) of vegetation types. For the USGS (Cogan et slides/slumps/flows and (3) mass movement/ al. 2004) data set we averaged affinity values colluvium/alluvial pediment mantle deposits across all matching species recorded in plots, (Table 1, Appendix 2). then further averaged across plots within NVC Environmental affinities of coalition groups associations. NVC associations sharing similar (relationships between plot-averaged species averaged affinities across all (10) coalition group affinities and environmental variables) groups were then identified through cluster were modeled using regression tree analysis analysis, using Pearson’s correlation coefficient (Breiman 1993, Crawley 2007) in SPLUS and average linkage hierarchical clustering in (Insightful Corp. 2005) via the tree function R (Crawley 2007, R Development Core Team with consistent splitting rules (mincut = 5, 2010). minsize = 10, mindev = 0.1) . Multiple runs of random 10-fold cross-validation were applied RESULTS to each model to determine the best number Coalition Groups and Their of terminal nodes minimizing total deviance Environmental Affinities (Crawley 2007), and models were pruned accordingly. Eleven coalition groups were identified through coalition clustering at a threshold Relating Groups to Previously-described value of phi = 0.15. One group containing 6 Vegetation Types species (Artemisia dracunculus, Elymus spica- We quantified the degree to which vegeta- tus, Sisymbrium altissimum, Galium aparine, tion types described by Harper et al. (2001) Physalis hederifolia, and Tragopogon dubius) and Cogan et al. (2004) were compositionally was excluded from further analysis because of similar to our coalition groups. This was ac- its limited distribution in disturbed low-eleva- complished by averaging, for each coalition tion environments. The remaining 10 groups group, species affinity values of matching contained 9–36 species each and collectively species attributed to each vegetation type. captured 169 of the 511 species recorded by 2015] PLANT COALITION GROUPS OF ZION NP 33 the Harper vegetation survey (Table 2, Appen- Crevice Canyon group had higher affinity dix 3). These coalition groups can be readily for higher elevations (Fig. 2C), exhibiting an described in terms of their environmental af - affinity pattern very similar to that of the finities at Zion, and we have given each group High Plateau group (Fig. 2B). The canopy a name that describes the environmental set- dominant species Abies concolor and the herbs ting for which it demonstrates greatest affinity Thalictrum fendleri and Taraxacum officinale (Table 2, Figs. 2–5). Coalition clustering’s al- were coalition species of both Crevice Canyon lowance for overlapping group membership and High Plateau (Table 2B–C). In addition, resulted in 34 cases of species belonging to 2 the Crevice Canyon group shared 4 species groups and one case of a species (Amelanchier (Acer negundo, Elymus canadensis, Equisetum utahensis) belonging to 3 groups. Environ- hymenale, and Agrostis exarata) with the mental affinities of coalition groups also over- Streambank group (Table 2A) and 5 (Heuchera lap to varying degrees, as depicted in predic- rubescens, Zauschneria latifolia, Holodiscus tive maps accompanying regression tree mod- dumosus, Selaginella underwoodii, Brickellia els (Figs. 2–5). In the group descriptions that grandiflora, and Erigeron sionis) with the follow, we highlight these cases of composi- Slickrock group (Table 2D). tional and environmental overlap and inter- As their names suggest, the Slickrock and pret their ecological significance. Upland Sands groups were differentiated by Three coalition groups were linked to mesic substrate preferences despite their common conditions at Zion: Streambank, High Plateau, affinity for Navajo Sandstone (Fig. 3). While and Crevice Canyon. The Streambank group having 4 coalition species in common (the (Table 2A) contained several species, including tree dominant Pinus ponderosa plus Arenaria Baccharis emoryi, Salix exigua, Tamarix chi- fendleri, Chrysopsis villosa, and Linanthas- nensis, and Populus fremontii, with affinities trum nuttallii) (Table 2D–E), the Slickrock for low-elevation riparian zones at Zion. Re- group was characterized by species typical of gression tree analysis confirmed the riparian exposed sandstone outcrops (e.g., Petrophytum setting: affinities were highest on alluvium or caespitosum, Cercocarpus intricatus, Castilleja lacustrine substrates below 1356 m with low scabrida, Muhlenbergia thurberi, and Ivesia topographic positions (TPI at 50-m radius < sabulosa) (Table 2D), whereas Upland Sands –0.51) (Fig. 2A). The Streambank group also species included the shrub dominant Arc- had moderately high affinities for low topo- tostaphylos patula plus several sand-loving graphic positions on other substrates (likely species (e.g., Cryptantha cinerea, Abronia fra- intermittent streams) and at high elevations grans, Artemisia campestris, Penstemon laevis, (>2321 m), presumably because mesic condi- and Tradescantia occidentalis) (Table 2E). The tions at Zion’s highest elevations are favorable high affinity of Upland Sands for Navajo Sand- for some riparian species. The High Plateau stone can be attributed to the group’s pres- group, which had highest affinity at elevations ence on pockets of sand smaller than the above 2243 m (Fig. 2B), shared 3 coalition mapping scale of the geologic layer (O’Meara species (Bromus carinatus, Elymus trachy- 2006). Plot data support this observation: in caulus, and Poa pratensis) with Streambank plots mapped to Navajo Sandstone, Slickrock (Table 2A–B). Coalition species unique to the affinity was positively correlated with percent High Plateau group include Stellaria jame- rock cover (Spearman’s rho = 0.50), while siana, Lupinus sericeus, Prunus virginiana, Upland Sands affinity was negatively corre- Stipa lettermanii, Rosa woodsii, and Juniperus lated (Spearman’s rho = –0.15). The Upland scopulorum. (Table 2B). Sands group also had relatively high affinity The Crevice Canyon group (Table 2C, Fig. for eolian sand deposits large enough to have 2C) occupied an environmental setting inter- been mapped by O’Meara (2006), as well as mediate between Streambank and High Pla - the sandstone-dominated Temple Cap for- teau: mid-elevation Navajo Sandstone canyons mation. Both Upland Sands and Slickrock where topographic shading and intermittent had higher affinities for higher elevations water flow produce cool mesic conditions. (>1753–1782 m) within the Navajo Sandstone Sites with these conditions have extremely zone (Fig. 3). Beyond the Navajo Sand - low TPI values (i.e., TPI at 50-m radius < stone, the Slickrock group had its highest –35; Fig. 2C). Outside of these canyons, the affinity value in sites with low TPI (TPI at 34 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

TABLE 2. Coalition and affiliate species of Zion National Park based on coalition clustering of 1987–1989 vegetation survey data. Occ. = number of plot occurrences out of 288 total. Affin. = affinity of a coalition species to its coalition group, calculated as the mean value of the phi coefficient of association between the species and others in the group. Affil. = affiliation of an affiliate species with coalition species, calculated as percent co-occurrence. Cvr. = mean cover of affiliate species weighted by co-occurrences with coalition species. Species in boldface are both coalition species and affiliate species for a given group. Underlined species are dominant affiliates (weighted mean cover ≥ 2%). Coalition species Occ. Affin. Affiliate species Occ. Affil. Cvr.

A. STREAMBANK Baccharis salicina 30.46Acer negundo 18 0.53 2.3 Salix exigua 30.46Bromus tectorum 130 0.51 5.1 Tamarix chinensis 50.43Bromus diandrus 10 0.47 8.3 Populus fremontii 60.39Equisetum hyemale 8 0.44 0.8 Equisetum hyemale 80.37Poa pratensis 14 0.42 4.3 Verbascum thapsus 50.37Artemisia ludoviciana 51 0.42 0.4 Sonchus sp. 30.37Phacelia heterophylla 42 0.41 0.3 Agrostis exarata 30.35Poa fendleriana 190 0.41 1.8 Bromus diandrus 10 0.34 Populus fremontii 6 0.40 4.6 Juncus arcticus 30.31Elymus trachycaulus 10 0.40 0.8 Elymus canadensis 40.27Chrysopsis villosa 74 0.39 1.6 Acer negundo 18 0.25 Tamarix chinensis 5 0.37 0.4 Bromus carinatus 90.25Bromus carinatus 9 0.36 0.6 Poa compressa 30.24Verbascum thapsus 5 0.35 0.2 Elymus trachycaulus 10 0.24 Quercus gambelii 135 0.34 5.7 Poa pratensis 14 0.20 Baccharis salicina 3 0.30 2.3 Datura wrightii 30.20Salix exigua 3 0.30 5.0 Fraxinus velutina 60.19Elymus canadensis 4 0.29 0.1 Gutierrezia sp. 94 0.28 0.3 Sonchus sp. 3 0.25 0.1 Agrostis exarata 3 0.24 0.1 Fraxinus velutina 6 0.23 4.2 Artemisia sp. 2 0.23 0.4 Melilotus sp. 2 0.23 0.1 Oenothera longissima 2 0.23 0.1 B. HIGH PLATEAU Stellaria jamesiana 18 0.33 Quercus gambelii 135 0.81 22.8 Senecio eremophilus 30.31Symphoricarpos oreophilus 58 0.65 11.0 Lupinus sericeus 90.29Vicia americana 43 0.61 5.1 Prunus virginiana 10 0.29 Poa fendleriana 190 0.61 2.3 Mertensia arizonica 70.29Polygonum douglasii 39 0.45 0.3 Bromus carinatus 90.27Amelanchier utahensis 141 0.44 3.1 Amelanchier alnifolia 11 0.27 Carex rossii 67 0.43 0.4 Stipa nelsonii 50.26Stellaria jamesiana 18 0.42 2.4 Stipa lettermanii 12 0.25 Arabis perennans 124 0.35 0.2 Vicia americana 43 0.25 Senecio multilobatus 99 0.32 0.2 Poa pratensis 14 0.25 Juniperus scopulorum 21 0.31 0.9 Rosa woodsii 12 0.24 Abies concolor 19 0.31 3.4 Achillea millefolium 60.20Phacelia heterophylla 42 0.29 0.2 Taraxacum officinale 11 0.20 Poa pratensis 14 0.29 4.4 Symphoricarpos oreophilus 58 0.20 Eriogonum racemosum 59 0.28 0.1 Thalictrum fendleri 12 0.20 Solidago velutina 38 0.27 1.2 Chenopodium fremontii 30.20Prunus virginiana 10 0.27 0.7 Abies concolor 19 0.19 Amelanchier alnifolia 11 0.27 0.7 Elymus trachycaulus 10 0.18 Stipa lettermanii 12 0.27 0.3 Juniperus scopulorum 21 0.18 Acer grandidentatum 29 0.26 2.7 Polygonum douglasii 39 0.17 Rosa woodsii 12 0.26 0.7 Osmorhiza occidentalis 40.16Opuntia macrorhiza 120 0.26 0.4 Bromus anomalus 50.16Pachystima myrsinites 54 0.26 1.6 Erigeron speciosus 60.15 C. CREVICE CANYON Eupatorium herbaceum 12 0.36 Quercus gambelii 135 0.69 7.1 Viola canadensis 50.33Poa fendleriana 190 0.67 4.5 Cystopteris fragilis 60.32Acer grandidentatum 29 0.60 6.2 Acer negundo 18 0.32 Acer negundo 18 0.52 5.5 Polypodium hesperium 60.27Eupatorium herbaceum 12 0.48 0.2 Galium triflorum 40.26Pachystima myrsinites 54 0.46 3.3 2015] PLANT COALITION GROUPS OF ZION NP 35

TABLE 2. Continued. Coalition species Occ. Affin. Affiliate species Occ. Affil. Cvr. Acer grandidentatum 29 0.26 Pseudotsuga menziesii 18 0.45 4.1 Rubus leucodermis 40.26Holodiscus dumosus 21 0.44 0.9 Bromus ciliatus 12 0.26 Solidago velutina 38 0.44 0.3 Pseudotsuga menziesii 18 0.26 Abies concolor 19 0.44 5.5 Heuchera rubescens 12 0.25 Draba asprella 33 0.39 0.2 Smilacina racemosa 60.25Artemisia ludoviciana 51 0.39 0.4 Mimulus guttatus 30.25Heuchera rubescens 12 0.38 0.2 Taraxacum officinale 11 0.24 Arabis perennans 124 0.38 0.2 Philadelphus microphyllus 10 0.24 Bromus ciliatus 12 0.38 0.2 Disporum trachycarpum 40.23Stephanomeria tenuifolia 51 0.37 0.2 Thalictrum fendleri 12 0.23 Zauschneria latifolia 15 0.37 0.2 Elymus canadensis 40.23Selaginella underwoodii 17 0.36 0.7 Abies concolor 19 0.22 Eriogonum racemosum 59 0.36 0.2 Epilobium glandulosum 30.22Chrysopsis villosa 74 0.35 0.7 Goodyera oblongifolia 50.21Penstemon rostriflorus 30 0.33 0.2 Muhlenbergia racemosa 30.21Senecio multilobatus 99 0.33 0.2 Aralia racemosa 30.20Philadelphus microphyllus 10 0.33 0.7 Sphaeromeria ruthiae 30.20Thalictrum fendleri 12 0.33 0.4 Zauschneria latifolia 15 0.20 Mahonia repens 29 0.33 0.5 Holodiscus dumosus 21 0.19 Taraxacum officinale 11 0.29 0.1 Chimaphila menziesii 50.19Phacelia heterophylla 42 0.29 0.3 Epilobium ciliatum 30.19 Epilobium brachycarpum 30.18 Selaginella underwoodii 17 0.17 Brickellia grandiflora 10 0.17 Erigeron sionis 40.17 Aster welshii 60.16 Pteridium aquilinum 30.16 Equisetum hyemale 80.16 Agrostis exarata 30.16 D. SLICKROCK Petrophytum caespitosum 12 0.31 Poa fendleriana 190 0.86 4.1 Holodiscus dumosus 21 0.31 Chrysopsis villosa 74 0.61 1.1 Cercocarpus intricatus 40 0.30 Quercus gambelii 135 0.56 5.6 Castilleja scabrida 40 0.29 Senecio multilobatus 99 0.56 0.3 Muhlenbergia thurberi 12 0.28 Pinus ponderosa 60 0.53 5.3 Selaginella underwoodii 17 0.27 Opuntia macrorhiza 120 0.53 0.5 Ivesia sabulosa 70.27Cercocarpus intricatus 40 0.51 1.5 Zauschneria latifolia 15 0.25 Stephanomeria tenuifolia 51 0.50 0.4 Arenaria fendleri 23 0.24 Castilleja scabrida 40 0.49 0.2 Erigeron canaani 16 0.23 Amelanchier utahensis 141 0.49 1.7 Eriogonum jamesii 10 0.23 Arabis perennans 124 0.48 0.2 Erigeron sionis 40.22Arctostaphylos patula 94 0.47 5.5 Leptodactylon pungens 80.22Arenaria macradenia 46 0.41 0.5 Stephanomeria tenuifolia 51 0.21 Carex rossii 67 0.40 0.3 Draba asprella 33 0.21 Solidago velutina 38 0.37 0.3 Trifolium longipes 10 0.20 Draba asprella 33 0.36 0.2 Penstemon humilis 27 0.20 Holodiscus dumosus 21 0.34 0.6 Pinus ponderosa 60 0.20 Eriogonum racemosum 59 0.32 0.2 Brickellia grandiflora 10 0.20 Penstemon humilis 27 0.32 0.2 Chrysopsis villosa 74 0.20 Arenaria fendleri 23 0.31 0.2 Monardella odoratissima 60.20Chaenactis douglasii 44 0.31 0.1 Solidago velutina 38 0.19 Phlox austromontana 54 0.30 0.4 Linanthastrum nuttallii 19 0.18 Pachystima myrsinites 54 0.28 1.4 Sedum lanceolatum 30.18Selaginella underwoodii 17 0.28 0.9 Heuchera rubescens 12 0.17 Bromus tectorum 130 0.27 0.6 Wyethia arizonica 50.17 Arenaria macradenia 46 0.15 E. UPLAND SANDS Cryptantha cinerea 17 0.26 Poa fendleriana 190 0.83 5.1 Chaenactis douglasii 44 0.25 Opuntia macrorhiza 120 0.72 1.1 Abronia fragrans 20 0.25 Senecio multilobatus 99 0.64 0.4 Hymenopappus filifolius 11 0.24 Arctostaphylos patula 94 0.62 10.1 36 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

TABLE 2. Continued. Coalition species Occ. Affin. Affiliate species Occ. Affil. Cvr. Artemisia campestris 14 0.23 Amelanchier utahensis 141 0.55 2.3 Penstemon laevis 26 0.23 Quercus gambelii 135 0.52 6.6 Tradescantia occidentalis 22 0.22 Chrysopsis villosa 74 0.50 1.5 Arctostaphylos patula 94 0.21 Arabis perennans 124 0.48 0.2 Opuntia macrorhiza 120 0.21 Erysimum asperum 64 0.43 0.2 Eriogonum alatum 60.20Pinus ponderosa 60 0.41 5.6 Chrysopsis villosa 74 0.19 Phlox austromontana 54 0.41 0.6 Senecio multilobatus 99 0.19 Bromus tectorum 130 0.39 1.1 Bouteloua gracilis 19 0.19 Chaenactis douglasii 44 0.39 0.2 Phlox austromontana 54 0.19 Juniperus osteosperma 115 0.36 2.2 Arenaria fendleri 23 0.19 Gilia inconspicua 84 0.34 0.2 Linanthastrum nuttallii 19 0.18 Festuca octoflora 74 0.33 0.3 Pinus ponderosa 60 0.17 Eriogonum racemosum 59 0.31 0.2 Erysimum asperum 64 0.16 Stephanomeria tenuifolia 51 0.30 0.2 Polygonum douglasii 39 0.15 Carex rossii 67 0.29 0.2 Polygonum douglasii 39 0.28 0.1 Sporobolus cryptandrus 50 0.27 0.2 Arenaria macradenia 46 0.27 0.3 Yucca angustissima 41 0.26 0.3 Penstemon laevis 26 0.25 0.2 Machaeranthera canescens 40 0.25 0.1 Stipa hymenoides 65 0.25 0.2 F. ROCKY SLOPES Fraxinus anomala 45 0.24 Poa fendleriana 190 0.85 5.8 Arabis perennans 124 0.24 Amelanchier utahensis 141 0.67 3.5 Erigeron utahensis 41 0.22 Arabis perennans 124 0.65 0.3 Poa fendleriana 190 0.21 Opuntia macrorhiza 120 0.61 1.0 Quercus turbinella 50 0.21 Juniperus osteosperma 115 0.58 4.1 Erysimum asperum 64 0.21 Senecio multilobatus 99 0.51 0.3 Opuntia macrorhiza 120 0.20 Bromus tectorum 130 0.50 1.3 Pinus monophylla 64 0.20 Quercus gambelii 135 0.46 7.0 Juniperus osteosperma 115 0.19 Gilia inconspicua 84 0.43 0.2 Haplopappus scopulorum 42 0.18 Arctostaphylos patula 94 0.42 6.2 Shepherdia rotundifolia 17 0.18 Erysimum asperum 64 0.38 0.2 Penstemon eatonii 23 0.18 Pinus monophylla 64 0.37 2.9 Amelanchier utahensis 141 0.17 Gutierrezia sp. 94 0.37 0.5 Senecio multilobatus 99 0.16 Quercus turbinella 50 0.32 3.6 Gilia inconspicua 84 0.15 Fraxinus anomala 45 0.30 0.9 Streptanthus cordatus 38 0.15 Festuca octoflora 74 0.30 0.2 Cryptantha sp. (annual) 60 0.29 0.2 Erigeron utahensis 41 0.28 0.2 Stipa hymenoides 65 0.28 0.2 Chrysopsis villosa 74 0.27 0.8 Haplopappus scopulorum 42 0.27 0.5 Eriogonum racemosum 59 0.27 0.1 Phlox austromontana 54 0.26 0.5 Pinus edulis 53 0.25 1.8 G. ARID LOWLANDS Bromus rubens 43 0.37 Bromus tectorum 130 0.78 3.4 Gutierrezia sp. 94 0.28 Gutierrezia sp. 94 0.70 1.3 Baileya multiradiata 80.27Juniperus osteosperma 115 0.69 4.1 Ephedra nevadensis 18 0.26 Gilia inconspicua 84 0.64 0.3 Draba verna 40 0.26 Poa fendleriana 190 0.61 3.2 Gilia inconspicua 84 0.26 Festuca octoflora 74 0.58 0.4 Hilaria sp. 41 0.25 Arabis perennans 124 0.49 0.3 Festuca octoflora 74 0.25 Cryptantha sp. (annual) 60 0.48 0.3 Coleogyne ramosissima 13 0.24 Bromus rubens 43 0.48 0.5 Opuntia erinacea 17 0.23 Pinus monophylla 64 0.45 2.6 Bouteloua eriopoda 50.23Opuntia macrorhiza 120 0.45 0.7 Eriogonum palmerianum 15 0.22 Draba verna 40 0.40 0.2 Astragalus nuttallianus 90.22Amelanchier utahensis 141 0.39 2.0 Bromus tectorum 130 0.21 Hilaria sp. 41 0.38 1.1 Cryptantha sp. (annual) 60 0.21 Stipa hymenoides 65 0.33 0.3 2015] PLANT COALITION GROUPS OF ZION NP 37

TABLE 2. Continued. Coalition species Occ. Affin. Affiliate species Occ. Affil. Cvr. Eriastrum eremicum 70.21Quercus turbinella 50 0.31 3.5 Psorothamnus fremontii 11 0.20 Elymus elymoides 58 0.30 0.2 Encelia frutescens 40.20Erigeron utahensis 41 0.28 0.2 Yucca baccata 17 0.19 Arctostaphylos patula 94 0.28 4.0 Stipa speciosa 15 0.19 Descurainia pinnata 28 0.25 0.2 Eriastrum sparsiflorum 13 0.18 Artemisia tridentata 55 0.24 1.7 Swertia albomarginata 11 0.18 Stipa comata 39 0.24 1.1 Descurainia pinnata 28 0.17 Eriogonum davidsonii 32 0.24 0.1 Rhus aromatica 23 0.17 Senecio multilobatus 99 0.23 0.1 Juniperus osteosperma 115 0.17 Eriogonum inflatum 30.17 Linanthus dichotomus 14 0.16 Pinus monophylla 64 0.16 H. LOWLAND FLATS Lycium pallidum 30.34Bromus tectorum 130 0.76 18.0 Erodium cicutarium 10 0.29 Gutierrezia sp. 94 0.58 2.0 Atriplex canescens 10 0.27 Chamaesyce albomarginata 23 0.54 1.2 Tragopogon dubius 80.24Juniperus osteosperma 115 0.48 1.6 Lactuca serriola 13 0.22 Sphaeralcea grossulariifolia 14 0.39 0.2 Cirsium wheeleri 80.22Erodium cicutarium 10 0.38 0.8 Chamaesyce albomarginata 23 0.21 Lactuca serriola 13 0.38 0.2 Sphaeralcea grossulariifolia 14 0.21 Hilaria sp. 41 0.36 1.3 Elymus smithii 30.19Atriplex canescens 10 0.35 0.9 Poa fendleriana 190 0.32 1.3 Sporobolus cryptandrus 50 0.30 0.5 Tragopogon dubius 8 0.28 0.1 Festuca octoflora 74 0.28 0.2 Stipa comata 39 0.28 1.4 Gilia inconspicua 84 0.27 0.1 Cirsium wheeleri 8 0.26 0.2 Amelanchier utahensis 141 0.26 1.5 Elymus elymoides 58 0.24 0.1 Quercus gambelii 135 0.24 2.6 Bromus rubens 43 0.23 0.3 Artemisia tridentata 55 0.22 1.8 Cryptantha sp. (annual) 60 0.22 0.1 I. UPPER MESA TOP Lathyrus lanszwertii 20 0.24 Quercus gambelii 135 0.78 15.1 Peraphyllum ramosissimum 10 0.24 Amelanchier utahensis 141 0.78 4.3 Balsamorhiza sagittata 80.22Poa fendleriana 190 0.78 4.3 Phlox longifolia 11 0.22 Arabis perennans 124 0.49 0.2 Penstemon linarioides 13 0.21 Opuntia macrorhiza 120 0.45 0.8 Swertia radiata 11 0.20 Carex rossii 67 0.45 0.5 Zigadenus paniculatus 12 0.19 Senecio multilobatus 99 0.43 0.2 Purshia tridentata 36 0.18 Juniperus osteosperma 115 0.43 2.5 Quercus gambelii 135 0.18 Symphoricarpos oreophilus 58 0.41 3.3 Symphoricarpos oreophilus 58 0.17 Pinus edulis 53 0.39 3.6 Mertensia fusiformis 30.17Arctostaphylos patula 94 0.38 5.3 Chrysothamnus depressus 50.16Pachystima myrsinites 54 0.33 1.5 Amelanchier utahensis 141 0.16 Pinus ponderosa 60 0.28 4.0 Carex rossii 67 0.16 Erysimum asperum 64 0.27 0.1 Pinus edulis 53 0.15 Bromus tectorum 130 0.27 0.8 Vicia americana 43 0.26 1.5 Phlox austromontana 54 0.25 0.7 Eriogonum racemosum 59 0.24 0.1 Cercocarpus montanus 33 0.23 2.0 Lathyrus lanszwertii 20 0.22 0.9 Solidago velutina 38 0.20 0.4 J. LOWER MESA TOP Viola purpurea 10 0.27 Amelanchier utahensis 141 0.80 4.2 Cercocarpus montanus 33 0.26 Quercus gambelii 135 0.77 13.9 Pinus edulis 53 0.22 Poa fendleriana 190 0.75 4.1 38 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

TABLE 2. Continued. Coalition species Occ. Affin. Affiliate species Occ. Affil. Cvr. Carex rossii 67 0.21 Arabis perennans 124 0.50 0.2 Pachystima myrsinites 54 0.21 Carex rossii 67 0.48 0.6 Amelanchier utahensis 141 0.20 Opuntia macrorhiza 120 0.44 0.8 Swertia radiata 11 0.19 Senecio multilobatus 99 0.44 0.2 Pedicularis centranthera 10 0.18 Pachystima myrsinites 54 0.43 1.8 Peraphyllum ramosissimum 10 0.18 Arctostaphylos patula 94 0.41 5.5 Quercus gambelii 135 0.18 Pinus edulis 53 0.41 4.0 Physaria newberryi 70.16Juniperus osteosperma 115 0.41 2.6 Mahonia repens 29 0.16 Symphoricarpos oreophilus 58 0.33 2.4 Cercocarpus montanus 33 0.31 2.9 Pinus ponderosa 60 0.27 3.6 Bromus tectorum 130 0.25 0.8 Erysimum asperum 64 0.25 0.1 Phlox austromontana 54 0.24 0.6 Vicia americana 43 0.24 1.2 Mahonia repens 29 0.24 0.3 Eriogonum racemosum 59 0.22 0.1 Fraxinus anomala 45 0.22 0.6

100-m radius > –21.24) that corresponded to multilobatus, and Erysimum asperum) with the talus deposits immediately below Navajo Sand- Upland Sands group (Table 2E) and 3 others stone cliffs (Fig. 3A). (Pinus monophylla, Juniperus osteosperma, and The group with highest affinity for the Gilia inconspicua) with the lower-elevation steep sloping terrain below Navajo Sandstone Arid Lowlands group (Table 2G). Pinus mono- cliffs was given the name “Rocky Slopes.” phylla and J. osteosperma are well-known domi - Fraxinus anomala, Erigeron utahensis, Poa nants of Great Basin woodlands (West 1999) fendleriana, and Quercus turbinella were and epitomize the Great Basin affinities of prominent members of this group (Table 2F). many of the Arid Lowlands coalition species. The first split of the Rocky Slopes regression Other coalition species of the Arid Lowlands tree model identified substrate as an impor- group, including Coleogyne ramossima, Bai- tant affinity predictor, with lower affinity for leya multiradiata, and Eriastrum eremicum, Carmel, alluvium, colluvium, eolian, lacus- have geographic affinities for the Mojave Desert trine and residuum than for other substrates and lower-elevation Colorado Plateau rather (Fig. 4A). Rocky Slopes affinity values were than the Great Basin (Welsh et al. 2003). In- low in the deep canyon zone (TPI at 100-m cluded in this mixture is the common Mojave radius < –35) (Fig. 4A) in contrast to the Desert invasive species Bromus rubens as well Crevice Canyon group (Fig. 2C), but higher as its Great Basin counterpart Bromus tectorum. in other low TPI areas adjacent to cliffs. The Gutierrezia spp. (G. sarothrae and G. micro- highest Rocky Slopes affinities were associ- cephala), indicators of grazing disturbance ated with the Kayenta formation which is par- history (Ladyman 2004), also had high affinity ticularly extensive in the Taylor Creek thrust for the Arid Lowlands group (Table 2G). fault area of northwestern Zion (Fig. 4A). The The arid character of sites occupied by the broad zone of relatively high affinity above, Arid Lowlands group was verified by its re - below, and around the Kayenta formation can gression tree model which indicated higher be attributed to the broad distribution of affinity for sites below 1596 m elevation (Fig. several members of the Rocky Slopes group, 4B). Within this low-elevation zone the Arid including Poa fendleriana, Amelanchier utah - Lowlands group had lower affinity for deep ensis, and Arabis perennans. These species canyons (TPI at 100-m radius < –22), sub- occupy a wide range of environments at Zion strates associated with riparian areas (alluvium but form a coalition with more narrowly dis- and lacustrine) and substrates concentrated at tributed species such as Shepherdia rotundifo- higher elevations (Kayenta, talus, Navajo and lia (Table 2F). eolian), except where these substrates were The Rocky Slopes group shared 3 coali- highly exposed (exposure > 215). At somewhat tion species (Opuntia macrorhiza, Senecio higher elevations (1596–1818 m) the Arid 2015] PLANT COALITION GROUPS OF ZION NP 39

Fig. 2. Regression tree models and maps for Streambank, High Plateau and Crevice Canyon coalition groups at Zion National Park, Utah. Mean affinity values at terminal nodes are in units of the phi coefficient. Predicted distribution of affinity values are mapped using shading shown at tree tips. TPI (50/100 m) = topographic position index (50- or 100-m radius). See Table 1 for key to substrate codes. 40 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

Fig. 3. Regression tree models and maps for Slickrock and Upland Sands coalition groups at Zion National Park, Utah. Mean affinity values at terminal nodes are in units of the phi coefficient. Predicted distribution of affinity values are mapped using shading shown at tree tips. TPI (100-m) = topographic position index at 100-m radius. See Table 1 for key to substrate codes.

Lowlands group likewise had higher affinity agriculture in the past and bear the marks for more exposed sites (exposure > 130) (Fig. of this land-use history. A combination of na - 4B) where the moisture/temperature regime tive species (e.g., Lycium pallidum, Atriplex was presumably similar to lower elevations. cane scens,Sphaeralcea grossulariifolia, Ely- Lowland Flats, another low-elevation species mus smithii) and exotics associated with dis- group, had high affinities for sites below turbance (Erodium circutarium, Tragopogon 1388 m, especially alluvium and Moenkopi dubius, Lactuca serriola) typified the Lowland substrates with TPI (50-m radius) > –0.5 Flats group (Table 2H). Various combinations (Fig. 4C). In Zion Canyon, these conditions of these species also occurred throughout Zion correspond with floodplains and benches at the lower elevations. Above 1388 m eleva- above the zone where the Streambank group tion, higher Lowland Flat affinities were as - is concentrated (Fig. 2A). These portions of sociated with flatter areas (slope < 6%) on Zion Canyon have been subject to intensive mostly unconsolidated substrates (Fig. 4C). 2015] PLANT COALITION GROUPS OF ZION NP 41

Fig. 4. Regression tree models and maps for Rocky Slopes, Arid Lowlands and Lowland Flats coalition groups at Zion National Park, Utah. Mean affinity values at terminal nodes are in units of the phi coefficient. Predicted distribu- tion of affinity values are mapped using shading shown at tree tips. TPI (50/100 m) = topographic position index (50- or 100-m radius). See Table 1 for key to substrate codes. 42 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

Fig. 5. Regression tree models and maps for Upper and Lower Mesa Top coalition groups at Zion National Park, Utah. Mean affinity values at terminal nodes are in units of the phi coefficient. Predicted distribution of affinity values are mapped using shading shown at tree tips. See Table 1 for key to substrate codes.

Coalition Clustering yielded 2 groups as - species (Pinus edulis, Quercus gambelii, Ame- sociated with the Carmel formation that caps lanchier utahensis, and Peraphyllum ramosissi- the Navajo Sandstone on the eastern and mum) as well as 2 herbaceous species (Carex northern sides of Zion. We interpreted one rossii and Swertia radiata), but the affinity group (Upper Mesa Top) as indicating slightly order of these species is almost exactly re - more mesic settings than the other group versed for the 2 groups, i.e., affinity of P. r a m o - (Lower Mesa Top). Highest affinity values for sissimum > S. radiata > A. utahensis > C. both groups were found in a zone defined by rossii > P. edulis for Upper Mesa Top (Table the Carmel formation above 1791 m, but out- 2I), whereas the opposite sequence is true for side of this zone the Upper Mesa Top group’s Lower Mesa Top (Table 2J). Two members of affinities are shifted toward higher elevations these groups overlap with other groups: Sym- than Lower Mesa Top’s (Fig. 5). The 2 Mesa phoricarpos oreophilus links Upper Mesa Top Top groups share a set of prominent woody to the High Plateau group (Table 2B) and 2015] PLANT COALITION GROUPS OF ZION NP 43

Amelanchier utahensis links both Mesa Top species were dominant in 5 or more groups groups to the Rocky Slopes group (Table 2F). (Quercus gambelii, Poa fendleriana, Ame- lanchier utahensis, Arctostaphylos patula, and Affiliate Species of Coalition Groups Juniperus osteosperma) (Table 2). Many but not all coalition species were Relations to Previously Described also affiliate species of their coalition groups Vegetation Types (Table 2, bold), meaning that they co-occurred with high frequency. The percentage of To a certain extent, our 10 coalition groups coalition species that were also affiliate resembled the 10 vegetation types previously species ranged from 36% for Crevice Canyon described by Harper et al. (2001). Some coali- group (Table 2C) to 83% for Streambank tion groups were compositionally very simi- (Table 2A). Each group also had 10–16 affiliate lar to modal species of Harper et al.’s (2001) species that were not coalition species (Table vegetation types (Fig. 6, Appendix 4). For ex - 2), meaning that they frequently co-occurred ample, 7 of the 9 coalition species of the Low- but did not share overall distribution/ land Flats group were modal species of the association patterns with members of the Abandoned Fields type (Appendix 4). The group. The total number of affiliate species, Arid Lowlands, Slickrock, High Plateau, and which reflects the mean species density in Streambank groups corresponded relatively plots where coalition species were concen- closely with the Blackbrush, Rock Crevice, trated, ranged from 21 for the Lower Mesa Douglas Fir–White Fir, and Riparian vege- Top group (Table 2J) to 27 for Crevice Canyon tation types, respectively (Fig. 6). However, (Table 2C). the correspondence was not exact, and some As a general pattern, the most widely dis- groups had high correspondence with multi- tributed species had the highest affiliation ple vegetation types, namely, Upland Sands values and were affiliated with the most coali- (with Ponderosa Pine, Mountain Brush, and tion groups. Poa fendleriana, the most fre- Rock Crevice), Crevice Canyon (with Ripar- quently encountered species at 190 occur- ian, Douglas/White Fir, and Hanging Gardens), rences, was an affiliate of all 10 described and the Mesa Top groups (with Douglas/ groups (Table 2). Other widely distributed spe- White Fir, Ponderosa Pine and Juniper-Pinyon) cies with multiple affiliations included Quer- (Fig. 6). cus gambelii, Amelanchier utahensis, Bromus When clustered according to coalition tectorum, Arabis perennans, Senecio multi - group affinities, NVC associations were ar - lobatus, and Opuntia macrorhiza (Table 2). In ranged into groups that reflected environ - each of these cases, the species was affiliated mental affinities but did not have uniform with several additional groups beyond the physiognomy (Fig. 7, Appendix 5). The 11 one(s) where it belonged as a coalition clusters highlighted in Fig. 7 were named species. Some species, including Artemisia for their predominant environmental (i.e., ludoviciana, Artemisia tridentata, Elymus coalition group) affinities and the mixture of elymoides, Eriogonum racemosum, Machaer- physiognomic classes they contained. These anthera canescens, Phacelia heterophylla, clusters were nested in 2 major groups (Xeric Sporobolus cryptandrus,Stipa comata, Stipa and Mesic Zion Vegetation) and 4 major sub- hymenoides, and Yucca angustissima, had groups (Fig. 7). Some associations emerged the distinction of being affiliate species with- as outliers whose environmental affinities out belonging to any of the coalition groups. seemed poorly matched with their neighbor- Weighted mean cover ranged from a frac- ing clusters (Needle-and-Thread Great Basin tion of a percent for most affiliate species Herbaceous, Coyote Willow/Barren Shrub- (Table 2) to 23% for Quercus gambelii in the land, and Green Rabbitbrush/Kentucky Blue- High Plateau group (Table 2B), followed by grass Semi-natural Shrubland) and these as - Bromus tectorum in the Lowland Flats group at sociations were excluded from the 10-cluster 18% (Table 2H). A total of 25 affiliate species classification (Fig. 7, Appendix 5). These out- met our criteria for dominance (weighted mean liers may be associations sampled by the cover > 2%) in relation to one or more groups USGS survey that were missed or poorly (Table 2, underlined entries). Ten of these captured by the Harper survey (see discus- were dominant in only one group, while 5 sion below). 44 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

Fig. 6. Correspondence between vegetation units of Harper et al. (2001) (rows) and coalition species groups of the current study (columns) at Zion National Park, Utah. Values shown are mean affinities (in units of the phi coefficient) of modal species of each vegetation unit in relation to each coalition group (see also Appendix 4).

DISCUSSION lack discrete spatial boundaries and reflect multispecies distribution patterns indepen- Our analysis of plant community variation dent of species commonness or abundance— and community-environment relationships at although we were also able to identify the Zion revealed several patterns that were over- most common and abundant species associ- looked or obscured in previous community ated with the coalition groups (affiliate and characterizations. Important patterns revealed dominant-affiliate species) in secondary analy- by our analysis include (A) floristic similarities sis steps. Our coalition groups likewise differ between communities of low-elevation ripar- from the NVC associations described by Cogan ian zones (Streambank) and high elevations et al. (2004), although the latter represent a (High Plateau), linked by mid-elevation com- finer scale of characterization that we were munities of narrow sandstone canyons (Crevice able to cross-walk to our broader coalition Canyon), (B) contrasting communities of sand- group scheme. stone bedrock (Slickrock) and sand deposits (Upland Sands) on the Navajo Sandstone For- Species Groups and the mation, (C) subtle community differentiation Individualistic Concept related to elevation within the Carmel Forma- Coalition groups can be considered species tion (Upper and Lower Mesa Top), and (D) groups according to most definitions of the continuity of communities containing Mojave term as used by plant community ecologists. Desert and Great Basin species within a xeric They are (phyto-) sociological species groups low-elevation zone (Arid Lowlands). comprised of species with similar association Our unique results can be attributed to the patterns (Doing 1969, Holzner et al. 1978) and way in which we defined and characterized also qualify as statistical species groups derived communities as coalition groups rather than through multivariate analysis (Bruelheide and conventional vegetation types. Although we Chytrý 2000, McCune and Grace 2002). Al- used the (nearly) identical dataset that Harper though our coalition groups are not ecological (1993) and Harper et al. (2001) used to charac- species groups as defined by European schools terize vegetation types at Zion, our characteri- of ecosystem classification (Mueller-Dombois zation was based on coalition clustering of and Ellenberg 1974, Kashian et al. 2003) they species association patterns as opposed to site are clearly “ecological” in the sense of being classification based on dominant species and correlated with environmental variation, such vegetation structure. Our 10 coalition groups that species within groups can be interpreted are comparable in scale to the 10 vegetation as having similar environmental affinities. types described by Harper et al. (2001) but Given the strength and interpretive value of 2015] PLANT COALITION GROUPS OF ZION NP 45

Fig. 7. Cluster dendrogram of National Vegetation Classification (NVC) associations of Zion National Park, Utah (Cogan et al. 2004), based on compositional affinities to 10 coalition groups presented in this paper. Names are assigned to clusters at 3 levels. A = Xeric Zion Vegetation, B = Mesic Zion Vegetation. A1 = Mesa Top/Upland Sands/Slickrock/ Rocky Slopes, A2 = Lowlands/Rocky Slopes/Upland Sands, B1 = High Plateau/Crevice Canyon, B2 = Streambank/ High Plateau. 46 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 these environmental correlations (Fig. 2–5), we tially identical to fuzzy membership, group- opted to use environmentally descriptive names averaged affinity to relative cardinality, and instead of following the widespread conven- plot-averaged affinity to fuzzy species richness tion of naming species groups based plant taxa as described by Olivero et al. (2011). (e.g. Holzner et al. 1978, Kashian et al. 2003). Although coalition clustering does not re - We propose that the term individualistic quire or produce spatially discrete community species group also applies to coalition groups units, it is influenced by discontinuities in because their nondiscrete spatial patterns, species distribution patterns. Multiple species overlapping composition and fuzzy attributes can have coinciding distributions marked by reflect notions of species individualism and quasi-discrete distribution boundaries due community continuity espoused by Gleason to abrupt environmental transitions, even if (1926, 1939). Gleason is well known for having these species would otherwise show unaligned pointed out difficulties of discrete community individualistic environmental responses (Aus- classification stemming from individualistic spe- tin and Smith 1989). Relatively abrupt transi- cies distributions, having noted that “every tions in hydrology, substrate and elevation are species of plant is a law unto itself, the distri - present at Zion and are reflected in our coali- bution of which in space depends upon its tion groups. The Streambank and Crevice individual peculiarities” (Gleason 1926, p. 26). Canyon groups, for example, were concen- But he also recognized that some species are trated in mesic environments immediately sufficiently aligned in their distributions to adjacent (at the scale of 10 × 10-m pixels) to warrant group recognition: xeric environments dominated by the Low- land Flats, Arid Lowlands or Slickrock groups The [individualistic] concept is by no means op - posed to the recognition of the synusia, or union, (Fig. 2–4). The Navajo Sandstone cliff zone defining it as a group of plants whose physiological forms a natural break in the Zion landscape demands are so similar that they are regularly se- that was detectable as a dividing line in re - lected by the same environment and consequently gression tree models, e.g., the first split of the regularly live together (Gleason 1939, p. 108). Arid Lowlands model at 1596 m elevation lies Hence an individualistic approach to com- at approximately the base of the Navajo Sand- munity characterization permits the dual objec- stone cliffs in the main section of the park tive of identifying spatially-associated species (Fig. 4B), while the first split of the Mesa Top possessing similar ecological requirements, groups (1791 m) lies near the top of this same while also recognizing and quantifying species set of cliffs (Fig. 5). differences. In other instances, our coalition clustering re - Unlike methods such as TWINSPAN or sults indicate gradual rather than abrupt changes COCKTAIL which characterize species groups in community composition and environmental in relation to discrete site groups (Bruelheide conditions. The 2 Mesa Top groups overlapped and Chytrý 2000), the coalition clustering extensively in composition (Table 2I–J), spatial approach extracts groups lacking discrete distribution and environmental affinities (Fig. boundaries (beyond the overall boundaries of 5), leading us to conclude that they represent the chosen study area). Any clustering method subtle distinctions along a continuum. The Arid carried out in R-mode (clustering species but Lowlands group also appears to represent a not sites) (McCune and Grace 2002) would continuum that was not subdivided at our se - likewise yield spatially nondiscrete groups. lected threshold level. We note, however, that Coalition groups have an added distinction of at a threshold of phi = 0.2 (instead of 0.15), 2 being compositionally nondiscrete (i.e., over- Arid Lowlands subgroups emerged, one with lapping in their species membership). The Great Basin/higher-elevation affinities and the only sense in which coalition groups are “dis- other with Mojave Desert/lower-elevation af - crete” is that any one group is composed of a finities (Fig. 8). Like the 2 Mesa Top groups, distinct set of member species, but even this these 2 Arid Lowlands subgroups had many form of discreteness is diluted by our use of species in common, and species with higher affinity values to characterize degrees of mem- affinity for one group tended to have lower af - bership. We note conceptual and mathematical finity for the other (with the exception of Bro- similarities between our affinity calculations mus rubens, which was the highest-affinity and fuzzy set theory: species affinity is essen- species for both groups) (Fig. 8). 2015] PLANT COALITION GROUPS OF ZION NP 47

did not actually co-occur in any plots, but the negative pairwise associations of these species were overridden by their shared positive asso- ciations with other species belonging to the group. Species occurring in multiple plots pro- vided a glue that held species occurring in fewer plots together in the same group. Our use of the phi coefficient contributed to this result because phi incorporates joint ab sence information and is less sensitive to differences in species frequency than presence-only coef- ficients such as the Jaccard index (Jackson et al. 1989, Sanderson et al. 2006). Species clus- tering based on presence-only coefficients tends to group species according to their fre- quency unless additional standardization steps are followed (Jackson et al. 1989, McCune and Grace 2002). The phi coefficient has the draw- Fig. 8. Two Arid Lowlands coalition groups at Zion National Park, Utah, obtained through coalition clustering back of being undefined when 2 species have in RCLUS at a threshold affinity value of phi = 0.2. perfectly overlapping distributions or when one Species are in descending order of group affinity and lines species’ distribution is nested within the other, connect species shared by both groups. The left group has but we did not encounter either of these situa- Great Basin (higher elevation) affinities and the right tions in our data set. Phi is always negative group has Mojave Desert (lower elevation) affinities. when 2 species never occur together, but can also be negative when 2 species occur together Coalition Clustering Methodology with few joint absences (e.g., a pair of ubiqui- We emphasize that our coalition groups tous species). In our analysis, co-occurring portray one of many possible coalition cluster- ubiquitous species would justifiably have been ing solutions, as illustrated by the alternative excluded from coalition groups, but this situa- Arid Lowlands groupings discussed above. tion was minimally relevant because the most Through experimentation with different asso- ubiquitous species (Poa fendleriana) occurred ciation coefficients and threshold levels, we in 66% of plots but no other species exceeded arrived at a solution at the desired scale that 50% occurrence. Our results would therefore agreed with our field experience at Zion. likely have been similar had we used a coeffi- Thus our approach differs from clustering cient that increases monotonically with in- approaches that seek to identify an optimal creasing joint presence, such as that of Baroni- set of clusters with minimal within-group and Urbani and Buser (1976). maximal between-group variation (Kaufman In contrast to Harper et al. (2001), whose and Rousseeuw 1990, Aho et al. 2008). Cluster community characterization began with domi- evaluation techniques (Aho et al. 2008) could nant species followed by prevalent and modal conceivably be applied to find an optimal species sensu Curtis (1959), our characteriza- threshold level resulting in a set of coalition tion began with a conceptual equivalent of groups with minimal species overlap. How- modal species (coalition species) followed by ever, given our assertion that overlapping equivalents of prevalent and dominant species composition contributes to the individualistic (affiliate and dominant-affiliate species, respec- character of coalition groups, we argue that tively). Modal species are conceptually similar flexibility in selecting a threshold level en- to what other authors have termed faithful, hances the usefulness of coalition clustering as diagnostic, differential or character species, all an exploratory tool. of which denote concentration of species occur- By applying coalition clustering to our data rences in one or more communities relative to using the phi coefficient of association we were others (Braun-Blanquet 1932, Aho et al. 2008, able to generate clusters containing mixtures Jennings et al. 2009). Coalition species like- of common and rarer species. In many cases, wise express this concept of community fidelity, species belonging to the same coalition group but in relation to a species group rather than 48 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 a set of spatially defined communities. Unlike teria by Harper et al. (2001) led to environ- strict definitions of faithful species, coalition mentally ambiguous vegetation types. Some species can be “faithful” to more than one group. of these ambiguities were clarified in a sub- Prevalent species as defined by Curtis (1959) sequent paper (Harper et al. 2003) that split are essentially synonymous with what others the Juniper-Pinyon vegetation type into 3 have termed constant species (Braun-Blanquet subtypes: a higher elevation subtype contain- 1932, Jennings et al. 2009), with the added cri- ing Pinus edulis (like the Mesa Top groups), a terion of using species density (average number mid-elevation subtype with Pinus monophylla of species per plot) to determine the number of (like Rocky Slopes) and a low-elevation sub- prevalent species for a community (Curtis 1959, type with Juniperus osteosperma but no Pinus Harper et al. 2001). Our procedures for identi- (like Arid Lowlands). Cogan et al. (2004) dif- fying affiliate and dominant-affiliate species ferentiated pinyon-juniper vegetation in a followed the logic of the prevalent species con- similar way at the NVC alliance level. cept in a way that, to our knowledge, is novel Our hierarchical rearrangement and reclas- and could be applied to any set of species sification of NVC associations on the basis of groups (coalition groups or otherwise). Although coalition group affinities (Fig. 7) differs sub- our measures of affiliation (percent co-occur- stantially from the way these associations have rence) and weighted mean cover are not directly been treated under the NVC system. Accord- comparable with constancy and mean cover ing to the NVC classification hierarchy fol- calculated for discrete communities, their values lowed by Cogan et al. (2004), associations were can be readily used for comparisons within nested within 6 major higher levels (alliance, and across species groups. formation, formation subgroup, formation group, formation subclass, formation class), each de - Perspectives on Vegetation Classification fined by specific characteristics including domi- By characterizing dominant species in rela- nant species, substrate, hydrologic regime, an - tion to species groups, we gained a different thropogenic disturbance, leaf phenology, leaf perspective of dominant species than previous structure, plant habit, and life form (Grossman authors who focused on them a priori for vege - et al. 1998). The NVC hierarchy was recently tation classification. Our analysis identified revised through the addition of new levels as dominant-affiliates many of the same species between alliance and formation (group, macro - that defined vegetation types of Harper et al. group and division) and changes to classifica- (2001), but some of these species were spread tion criteria designed to make groupings more among multiple coalition groups. Pinus pon- ecologically meaningful (FGDC 2008, Faber- derosa, which defined the Ponderosa Pine Langendoen et al. 2009, NatureServe 2013, vegetation type of Harper et al. (2001), emerged USNVC 2013). Upper levels of the new NVC as a dominant-affiliate species of 4 coalition hierarchy reflect general growth form differ- groups (Upper and Lower Mesa Top, Upland ences indicative of broad-scale environments Sands and Slickrock); likewise Abies concolor or “global macroclimatic factors”, while suc- of the White Fir–Douglas Fir vegetation type cessively lower levels reflect finer physiog- was a dominant-affiliate of both High Plateau nomic and floristic distinctions indicative of and Crevice Canyon; and dominants of Juniper- finer-scale climate, geology, substrate, hydrol- Pinyon (Pinus edulis, Pinus monophylla, Juni - ogy, moisture/nutrient factors, and disturbance perus osteosperma) and Mountain Brush (Quer- regimes (Faber-Langendoen et al. 2009). The cus gambelii, Amelanchier utahensis) vegetation NVC hierarchy might thus be expected to types were dominant-affiliates of 2 or more portray environmental affinities at Zion in a coalition groups each (Table 2). These mis- manner similar to our coalition groups; how- matches highlight the discordance between ever, our coalition group–based hierarchy was abundance patterns of single species and poorly aligned with both the older and presence-absence patterns of broader species newer NVC classification (Appendixes 6, 7). sets. Because dominant species and growth Some NVC groupings were concentrated within forms such as ponderosa pine, white fir, certain clusters of our hierarchy—e.g., Singleleaf juniper-pinyon and mountain brush are gen- Pinyon – (Utah Juniper) Woodland Alliance in eralists that occupy multiple environmental the Rocky Slopes Woodland/Shrub cluster—but settings at Zion, their use as classification cri- others were spread across multiple clusters, as 2015] PLANT COALITION GROUPS OF ZION NP 49 illustrated by the mixture of associations with but unlike the NVC and TESC it cannot be multiple physiognomies (forest, woodland, straightforwardly transferred to other land- shrubland, etc.) within groupings of our 3- scapes with different combinations of species tiered classification (Fig. 7). These differences and environments. can be attributed to the emphasis placed on Sampling Strategy and Modeling Issues physiognomy and dominant species at all levels of the NVC hierarchy, in contrast to our strictly Our ability to extrapolate coalition group floristic, presence/absence–based approach to affinities beyond park boundaries or interpo- classification. late them beyond plot locations within the Comparisons can also be made between park was limited by sampling and modeling our coalition group–based hierarchy of NVC constraints (cf. Elith and Leathwick 2009). associations (Fig. 7) and the Terrestrial Eco- Environmental conditions such as higher or logical Systems Classification (TESC) that has lower elevations in the surrounding region been applied at Zion (Comer et al. 2003, beyond the park were not sampled, and Comer and Schulz 2007, NatureServe 2013). important community-environment variation The TESC was developed to complement the within the park may have also been missed. NVC by grouping associations or alliances that Furthermore, the environmental variables we “tend to naturally co-occur with similar envi- used posed limitations on predictive model- ronmental settings, ecological dynamics, and/or ing; for example, the lacustrine substrate class environmental gradients” (Comer and Schulz noticeably overpredicted Streambank (Fig. 2A) 2007, p. 326), forming “ecological systems” and underpredicted Rocky Slopes (Fig. 4A) that can more easily be mapped and related affinities in the Coalpits Wash area (south- to environmental variation at landscape to re - western side of park), where—unlike in Zion gional scales (Comer et al. 2003, Comer and Canyon—lacustrine deposits are not associ- Schulz 2007). An ecological system can include ated with riparian areas. multiple physiognomic types that occur in Ideally, a different set of more natural land- a similar environment, such as successional scape boundaries could have been used to stages ranging from herbaceous vegetation to delimit our study area; but given that national shrubland to woodland within a riparian flood- park boundaries were used, the mostly sys- plain (Comer and Schulz 2007). Given these tematic sampling strategy followed by the criteria, the TESC approach could potentially Harper survey was advantageous for capturing yield ecological groupings similar to those we community and environmental variation in a identified at Zion. Upon cross-walking our relatively unbiased way. Dismissing the unlikely coalition-group based hierarchy with ecologi- possibility that the approximately 1.6-km dis- cal systems (Appendix 8), we did indeed find tance between Harper survey plots coincides general similarities, such as alignment of the with periodic environmental variation at Zion, North American Warm Desert Lower Montane the more-or-less evenly spaced plot arrange- Riparian Woodland and Shrubland system with ment (Fig. 1A) can be viewed as an area- the Streambank group. However, the concor- proportional sample of Zion’s environments. dance between the 2 classifications is far from Environmental conditions that were more exact, and we note that the ecological systems frequent in the landscape were captured with contain less physiognomic variability than clus- greater frequency by the Harper survey, in ters in our hierarchy (Fig. 7, Appendix 8). Like contrast to the USGS survey, which sought to the NVC hierarchy, the TESC as currently capture the range of environmental variation applied to vegetation types at Zion appears to without regard for frequency through a grad- emphasize physiognomic/structural uniformity sect sampling strategy (Cogan et al. 2004). The at the expense of environmental similarity. These effects of these different sampling strategies can observations are not intended as criticisms of be illustrated by histograms of plot-environ- the NVC and TESC approaches but rather ment combinations compared proportionally highlight their strengths and weaknesses in with background environmental variation at relation to our approach. No single commu- Zion (Figs. 9, 10) (Calenge 2006). The Harper nity characterization approach is likely to serve data set had an overrepresentation of low ele- all purposes; hence our characterization is vations, low slopes, moderate exposures (Fig. finely tuned to represent conditions at Zion, 9A), and alluvial substrates (Fig. 10A) that can 50 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

Fig. 9. Histograms showing background frequencies of topographic variables at Zion National Park, Utah (solid border bars), compared proportionally with plot representation of the same variables (shaded bars) by 2 different vegetation surveys (left and right). Based on 50-m-resolution generalizations of 10-m-resolution grids covering the park. Derived using function “histniche” of the adehabitat package (Calenge 2006) in R 2.8.0 (R Development Core Team 2008).

Fig. 10. Histograms showing background frequencies of geologic substrate variables at Zion National Park, Utah (solid border bars), compared proportionally with plot representation of the same variables (shaded bars) by 2 different vegetation surveys (left and right). See Table 1 for key to substrate codes. Based on 50-m-resolution generalizations of 10-m-resolution grids covering the park. Derived using function “histniche” of the adehabitat package (Calenge 2006) in R 2.8.0 (R Development Core Team 2008). be attributed at least in part to the nonsystem- underrepresentation were present to an even atic riparian and old field plots included in the greater degree in the USGS data set (Fig. 9B, survey. On the other hand, excessively steep 10B). Consequently the Harper data set has or bare plot locations were avoided or adjusted a larger, comparatively more representative by Harper survey workers, leading to an under- sample of common environments (such as the representation of high slopes and Navajo Sand- Navajo Sandstone), while the USGS data set stone substrates (Fig. 9A, 10A). However, these has better representation of rarer, more unique same patterns of environmental over- and environments and communities. The Harper 2015] PLANT COALITION GROUPS OF ZION NP 51 survey captured “transitional” areas with het- after fire could later die out, if for example erogeneous vegetation that would have been invasive Bromus species become dominant avoided by the USGS survey, while USGS and cause recurring fires (Bagchi et al. 2013). likely captured spatially restricted vegetation The recent fires at Zion could have therefore types that were missed by Harper. These dif- altered some of the community patterns we ferences between survey data sets, as well as have presented here, though this does not differences due to year of sampling, plot size, diminish the value of our characterization as and data collection techniques, must be kept a baseline reference. in mind when comparing our results with Long-term fire history has been implicated results derived from the USGS survey (i.e., as a factor determining current distributions Fig. 7, Appendixes 5–8). of pinyon-juniper woodland versus mountain brush in areas that could support either vege- Temporal Scale, Fire, and Weather tation type, with mountain brush occupying Our community characterization is best sites where the fire-return interval has been viewed as a snapshot that captures community shorter (Floyd et al. 2000). Although our coali- patterns in relation to environment conditions tion clustering analysis did not reveal discrete at a specific area within a specific timeframe. differences between these 2 vegetation types, Conditions at Zion in the late 1980s when the the 2 Mesa Top coalition groups were partially Harper survey was conducted included the ef - aligned with them. The fire-sensitive species fects of previous decades of agricultural dis- Pinus edulis and Cercocarpus montanus had turbance, livestock grazing, and fire suppres- higher affinity for the Lower Mesa Top group sion (Woodbury 1933, Madany and West 1983) than fire-tolerant mountain brush species but not the effects of more recent wildfires Quercus gambelii, Amelanchier utahensis, Bal- within the park, such as the 4256-ha Kolob samorhiza sagittata, and Symphoricarpos oreo- Fire of 2006 (Brisbin et al. 2013). Most Harper philus (Floyd et al. 2000, USDA–FS 2013), Survey plots were classified by field workers but in the Upper Mesa Top group the fire- as late-seral or climax with respect to time tolerant species had higher affinity (Table 2I–J). since fire, although 2 plots on the Horse Pas- These 2 Mesa Top groups might thus reflect ture Plateau burned in 1988 and were sam- differences in long-term fire history that have pled later the same year. These recently-burned left a subtle imprint on species presence- plots, despite their reduced cover relative to absence patterns. surrounding woodlands, readily blended into The patterns we detected at Zion also re - the Mesa Top coalition groups in our analy - flect weather conditions and fluctuations dur- sis, suggesting that the fire had a minimal effect ing the years leading up to the time of data on species composition as measured by pres- collection in 1987–1989. Weather data from ence-absence, at least initially. This supports the park show periods of above-average pre- the view that species presence-absence pat- cipitation during the years 1978–1983, near- terns are less sensitive to short-term environ- average from 1984–1988, and below-average mental fluctuations and disturbances than in 1989 (Witwicki 2013). Annual species sensi- abundance patterns, making presence-absence tive to yearly precipitation fluctuations may a more stable measure indicative of longer- have been sparser in 1989; however, most term ecological processes than abundance sampling of low elevations where most of these (Allen and Starr 1982, Wilson 2012). However, annual species occur took place in 1987–1988. the stability of presence-absence patterns can Since the late 1980s, Zion has experienced be expected to vary due to variation in plant large fluctuations in annual precipitation and longevity and sensitivity to disturbance. Other a gradual warming trend (Witwicki 2013) that studies in plant communities similar to those is projected to continue (Polley et al. 2013). In at Zion have reported shifts in species presence- light of evidence from elsewhere in south- absence due to fire, often characterized by western North America (Kelly and Goulden the loss of fire-intolerant shrubs and trees 2008, Fellows and Goulden 2012, Brusca et al. combined with an influx of annual species 2013), upward elevational shifts in plant species (Koniak 1985, Paysen et al. 2000, Ott et al. distributions could have occurred at Zion dur- 2001, Huisinga et al. 2005, Engel and Abella ing this period. Relative to other areas with 2011). Some species that might persist initially less abrupt elevation gradients, plant species 52 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 at Zion might be expected to easily disperse to Plateau. Pages 147–153 in E.D. McArthur and D.J. higher elevations, although species with affini- Fairbanks, compilers, Shrubland ecosystem genetics and biodiversity: proceedings. Proc. RMRS-P-21, ties for certain horizontally-layered geologic USDA Forest Service, Rocky Mountain Research substrates might find their preferred substrate Station, Ogden, UT. unavailable higher up. The degree to which BEYER, H.L. 2004. Hawth’s Analysis Tools for ArcGIS. coalition groups will remain associated through http://www.spatialecology.com/htools continued climate change is unknown, but BIEK, R.F., G.C. WILLIS, M.D. HYLLAND, H.H. DOELLING, D.A. SPRINKEL, T.C. CHIDSEY JR., AND P. B . A NDERSON. from a probabilistic perspective, we suggest 2 2003. Geology of Zion National Park, Utah. Pages general rules: (1) species will likely become 107–137 in Anonymous, editor, Geology of Utah’s increasingly disassociated with the passage of parks and monuments. Utah Geological Association time as their individualistic responses to cli- and Bryce Canyon Natural History Association. BRAUN-BLANQUET, J. 1932. Plant sociology: the study of mate change emerge, yet (2) species currently plant communities [English translation]. McGraw- more strongly associated will likely remain Hill, New York, NY. associated for longer than more weakly associ- BREIMAN, L. 1993. Classification and regression trees. ated species. We suggest that species associa- Chapman and Hall, New York, NY. BRISBIN, H., A. THODE, M. BROOKS, AND K. WEBER. 2013. tion patterns such as those we have quantified Soil seed bank responses to postfire herbicide and for Zion have predictive value in both space native seeding treatments designed to control Bro- and time. mus tectorum in a pinyon-juniper woodland at Zion National Park, USA. Invasive Plant Science and Management 6:118–129. CKNOWLEDGMENTS A BRUELHEIDE, H., AND M. CHYTRÝ. 2000. Towards unifica- tion of national vegetation classifications: a compari- We thank the many individuals mentioned son of two methods for analysis of large data sets. by Harper et al. (1992) and Cogan et al. (2004) Journal of Vegetation Science 11:295–306. who were involved in different stages of their BRUSCA, R.C., J.F. WIENS, W.M. MEYER, J. EBLE, K. FRANKLIN, J.T. OVERPECK, AND W. M OORE. 2013. Dra- respective vegetation surveys. The original matic response to climate change in the Southwest: Harper survey was partially funded by the U.S. Robert Whittaker’s 1963 Arizona mountain plant tran- National Park Service, and the work presented sect revisited. Ecology and Evolution 3:3307–3319. here was partially funded by the U.S. Forest [BLM] BUREAU OF LAND MANAGEMENT. 2006. Geographic Coordinate Data Base. http://www.blm.gov/wo/st/en/ Service, Rocky Mountain Research Station. Ad - prog/more/gcdb.html ditional support came from the University of CALENGE, C. 2006. 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Appendix 1 on page 55 Appendix 2 on page 55 Appendix 3 on page 56 Appendix 4 on page 75 Appendix 5 on page 80 Appendix 6 on page 84 Appendix 7 on page 90 Appendix 8 on page 96 2015] PLANT COALITION GROUPS OF ZION NP 55

APPENDIX 1. Abiotic environmental variable layers derived from a 10-m-resolution digital model of Zion National Park, Utah.

APPENDIX 2. Geologic substrate classes derived from a 1:24,000-scale digital geologic map of Zion National Park, Utah, and vicinity (O’Meara 2006). 56 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 bank bank 0.0410 0.0410 0.0090 0.0429 0.2513 0.0380 0.0817 0.1104 0.0533 0.0067 0.0094 0.0390 0.3453 0.1104 0.2268 0.0119 0.0006 0.0627 0.1007 0.1104 -0.0360 -0.0360 -0.0025 -0.0085 -0.0085 -0.0085 -0.1249 -0.0159 Stream- Stream- 0.2236 0.2236 0.1461 0.2614 0.3163 0.0272 0.2531 0.1540 0.2611 0.1570 0.2611 0.0093 0.0107 0.1498 0.0281 0.2611 Crevice -0.0349 -0.0349 -0.0098 -0.0012 -0.0098 -0.0098 -0.0012 -0.0269 -0.0098 -0.0098 -0.0817 -0.0299 Canyon Canyon High 0.1859 0.1859 0.0185 0.1139 0.0499 0.2048 0.0479 0.0132 0.0581 0.2240 0.0257 0.0870 0.0565 0.0416 0.0581 0.0068 0.0793 0.2711 0.0581 -0.0325 -0.0325 -0.0032 -0.0055 -0.0127 -0.0061 -0.0446 -0.0220 -0.0162 Plateau Plateau s to coalition species groups. # Occ. = Upper Upper 0.0229 0.0229 0.0003 0.0136 0.0053 0.0458 0.0051 0.0507 0.1598 0.0075 -0.0382 -0.0382 -0.0116 -0.0579 -0.0186 -0.0308 -0.0138 -0.0040 -0.0059 -0.0138 -0.0332 -0.0138 -0.0217 -0.0045 -0.0019 -0.0252 -0.0232 -0.0138 Mesa Top Top Mesa ient) between a species and members of coalition group. Lower Lower 0.0603 0.0603 0.0035 0.0312 0.0006 0.0140 0.0140 0.0570 0.1990 0.0140 -0.0448 -0.0448 -0.0447 -0.0178 -0.0352 -0.0150 -0.0051 -0.0150 -0.0033 -0.0150 -0.0207 -0.0248 -0.0161 -0.0033 -0.0024 -0.0282 -0.0177 -0.0414 Mesa Top Top Mesa 0.0865 0.0865 0.0395 0.1020 0.0587 0.0736 0.0046 0.0224 0.0370 0.0010 0.0083 0.0370 0.1228 0.1292 0.0231 0.0669 0.0370 -0.0028 -0.0028 -0.0163 -0.0163 -0.0056 -0.0099 -0.0163 -0.0213 -0.0029 -0.0081 -0.0266 Sands Slickrock Slickrock Sands 0.2521 0.2521 0.0190 0.0492 0.0599 0.0856 0.0303 0.0340 0.0390 Upland Upland -0.0559 -0.0559 -0.0307 -0.0489 -0.0678 -0.0189 -0.0340 -0.0027 -0.0149 -0.0149 -0.0094 -0.0073 -0.0112 -0.0211 -0.0027 -0.0240 -0.0447 -0.0027 -0.0027 Rocky Rocky Slopes Slopes 0.0253 0.0253 0.0044 0.1661 0.0268 -0.0770 -0.0770 -0.0117 -0.0451 -0.0929 -0.0386 -0.0467 -0.0466 -0.0291 -0.0368 -0.0067 -0.0290 -0.0142 -0.0469 -0.0291 -0.0329 -0.0317 -0.0064 -0.0216 -0.0014 -0.0809 -0.0358 -0.0291 Arid Arid 0.0051 0.0051 0.0230 -0.0893 -0.0893 -0.0051 -0.0484 -0.0988 -0.0711 -0.0443 -0.0290 -0.0290 -0.0204 -0.0133 -0.0071 -0.0144 -0.0204 -0.0164 -0.0204 -0.0183 -0.0403 -0.0156 -0.0204 -0.0075 -0.0668 -0.0968 -0.0164 -0.0204 Lowlands Lowlands Flats Flats 0.0196 0.0196 0.0537 0.0689 0.0062 0.0543 0.1286 -0.0226 -0.0226 -0.0448 -0.0269 -0.0556 -0.0226 -0.0003 -0.0154 -0.0154 -0.0109 -0.0109 -0.0109 -0.0189 -0.0109 -0.0109 -0.0113 -0.0109 -0.0109 -0.0245 -0.0900 -0.0109 Lowland Lowland 6 6 2 2 1 1 1 1 1 1 3 1 2 9 1 1 2 3 1 20 29 18 11 18 0.15 indicate membership in a group. Green shading by cell value is scaled separately for each column. Nomenclature follows 141 141 ≥ Nyctaginaceae Aceraceae Aceraceae Aceraceae Compositae Ranunculaceae Polypodiaceae Polypodiaceae Labiatae Compositae Compositae Compositae Gramineae Gramineae Gramineae Gramineae Liliaceae Liliaceae Liliaceae Liliaceae Rosaceae Rosaceae Gramineae Liliaceae Umbelliferae Pinaceae 19 Pinaceae 3. Vascular plant species recorded by the Harper vegetation survey (1987–1989) at Zion National Park, Utah, and their affinitie plant species recorded by the Harper vegetation survey (1987–1989) at Zion National Park, 3. Vascular

sp. PPENDIX A Scientific Name Name Scientific Abronia fragrans Acer glabrum Family Acer grandidentatum Acer negundo millefolium Achillea rubra Actaea # Occ. capillus-veneris Adiantum Adiantum pedatum Agastache urticifolia aurantiaca Agoseris glauca Agoseris retrorsa Agoseris cristatum* Agropyron exarata Agrostis scabra Agrostis stolonifera* Agrostis Allium acuminatum Allium macropetalum nevadense Allium Allium Amelanchier alnifolia utahensis Amelanchier gerardii Andropogon breviflorum Androstephium Angelica pinnata Abies concolor Abies number of plot occurrences out of 288 plots total. Affinities are mean pairwise association values (in units of the phi coeffic number of plot occurrences out 288 plots total. Affinities species that have at least 3 occurrences, values For et al. (2003). Welsh 2015] PLANT COALITION GROUPS OF ZION NP 57

bank bank tream- 0.0540 0.0540 0.0817 0.0469 0.0469 0.0050 0.0031 0.1026 0.4632 -0.0085 -0.0085 -0.0121 -0.0085 -0.0070 -0.0070 -0.0025 -0.0085 -0.0931 -0.0192 -0.0085 -0.0949 -0.0427 -0.0588 -0.0085 -0.0085 -0.0085 -0.0095 -0.0327 -0.0138 -0.0016 -0.0085 -0.0200 S

ce i 0.1447 0.1447 0.2110 0.0179 0.0585 0.0585 0.1668 0.2012 0.0963 0.0005 0.0962 0.0312 -0.0098 -0.0098 -0.0139 Crev -0.0098 -0.0098 -0.0301 -0.0220 -0.0016 -0.0837 -0.0203 -0.0037 -0.0098 -0.0098 -0.0393 -0.0098 -0.0315 -0.0260 -0.0170 -0.0016 -0.0710 Canyon Canyon h g i H 0.0635 0.0635 0.0476 0.0385 0.0385 0.0474 0.0313 0.0096 0.0214 0.1245 0.0580 0.0195 0.0197 -0.0127 -0.0127 -0.0136 -0.0127 -0.0163 -0.0163 -0.0499 -0.0202 -0.0959 -0.0512 -0.0401 -0.0127 -0.0058 -0.0646 -0.0004 -0.0341 -0.0225 -0.0220 -0.0021 Plateau Plateau

Upper 0.0148 0.0148 0.0428 0.0428 0.0240 0.0083 0.0110 0.0301 0.0034 0.0998 0.0354 -0.0138 -0.0138 -0.0054 -0.0030 -0.0217 -0.0232 -0.0232 -0.0238 -0.0151 -0.0124 -0.0522 -0.0069 -0.0124 -0.0138 -0.0888 -0.0361 -0.0178 -0.0308 -0.0201 -0.0308 -0.0307 Mesa Top Top Mesa

Lower 0.0195 0.0195 0.0152 0.0065 0.0065 0.0472 0.0106 0.0411 0.0893 0.0778 0.0007 0.0247 -0.0150 -0.0150 -0.0122 -0.0248 -0.0250 -0.0250 -0.0147 -0.0187 -0.0132 -0.0605 -0.0052 -0.0132 -0.0150 -0.0986 -0.0462 -0.0208 -0.0431 -0.0376 -0.0352 -0.0650 Mesa Top Top Mesa 0.0019 0.0019 0.0238 0.0270 0.0919 0.0919 0.0010 0.0178 0.0767 0.2407 0.1502 0.0148 0.0069 0.0346 -0.0035 -0.0035 -0.0163 -0.0109 -0.0109 -0.0017 -0.0054 -0.0047 -0.0109 -0.0163 -0.0026 -0.0041 -0.0146 -0.0044 -0.0291 -0.0250 -0.0196 -0.0941 d an l Sands Slickrock Slickrock Sands 0.0034 0.0034 0.0489 0.0173 0.0173 0.0123 0.2147 0.1880 0.0990 0.0579 0.2341 0.0261 0.0087 -0.0057 -0.0057 -0.0189 -0.0240 Up -0.0269 -0.0269 -0.0286 -0.0035 -0.0417 -0.0240 -0.0097 -0.0240 -0.0051 -0.0097 -0.0278 -0.0069 -0.0349 -0.0115 -0.0290

y k Roc 0.0049 0.0049 Slopes Slopes 0.0010 0.0010 0.2379 0.1003 0.1132 0.1271 0.0011 0.1161 0.0140 -0.0063 -0.0063 -0.0021 -0.0414 -0.0368 -0.0509 -0.0509 -0.0466 -0.0507 -0.0368 -0.0368 -0.0408 -0.0059 -0.0138 -0.0142 -0.0399 -0.0323 -0.0022 -0.0347 -0.0216 -0.0370 id id Ar 0.0083 0.0083 0.0295 0.0295 0.0189 0.1291 0.0497 0.0007 0.0237 -0.0159 -0.0159 -0.0276 -0.0290 -0.0204 -0.0355 -0.0355 -0.0290 -0.0204 -0.0331 -0.0204 -0.0183 -0.0204 -0.0593 -0.0513 -0.0607 -0.0001 -0.0204 -0.0411 -0.0471 -0.0227 -0.0204 -0.0198 Lowlands Lowlands d an l Flats Flats 0.0512 0.0512 0.0216 0.0383 0.0695 0.0211 -0.0109 -0.0109 -0.0154 -0.0189 -0.0154 -0.0109 -0.0189 -0.0189 -0.0154 -0.0109 -0.0953 -0.0189 -0.0109 -0.0245 -0.0109 -0.1013 -0.0544 -0.0804 -0.0109 -0.0109 -0.0334 -0.0189 -0.0109 -0.0089 -0.0154 Low 1 2 3 2 1 2 1 3 1 5 1 1 1 1 4 3 1 2 94 23 46 26 14 12 51 55 124 124 Compositae Compositae Compositae Compositae Apocynaceae Ranunculaceae Ranunculaceae Ranunculaceae Cruciferae Cruciferae Araliaceae Viscaceae Viscaceae Viscaceae Ericaceae Caryophyllaceae Caryophyllaceae Caryophyllaceae Papaveraceae Gramineae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Ranunculaceae 3 Ranunculaceae

3. Continued. sp. sp. PPENDIX A Antennaria microphylla Antennaria Antennaria parvifolia Antennaria rosulata androsaemifolium Apocynum chrysantha Aquilegia Aquilegia formosa formosa Aquilegia Aquilegia Arabis hirsuta Arabis perennans Aralia racemosa abietinum Arceuthobium divaricatum Arceuthobium vaginatum Arceuthobium patula Arctostaphylos fendleri Arenaria macradenia Arenaria pusilla Arenaria munita Argemone purpurea Aristida cordifolia Arnica arbuscula Artemisia campestris Artemisia dracunculus Artemisia Artemisia filifolia frigida Artemisia ludoviciana Artemisia Artemisia tridentata Artemisia Scientific Name Name Scientific Family # Occ. 58 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.4594 0.4594 0.1572 0.0360 0.0031 0.0533 0.1156 0.1156 0.0906 0.1134 0.0144 0.0003 0.0038 -0.0121 -0.0121 -0.0245 -0.0245 -0.0121 -0.0192 -0.0306 -0.0082 -0.0192 -0.0025 -0.0193 -0.0193 -0.0085 -0.0085 -0.0085 -0.0260 -0.0085 -0.0311 -0.0275 S

ce i 0.0332 0.0332 0.0117 0.0529 0.1723 0.0747 0.1089 0.1160 0.1160 0.0050 0.1633 Crev -0.0139 -0.0139 -0.0280 -0.0256 -0.0089 -0.0220 -0.0361 -0.0159 -0.0092 Canyon Canyon -0.0264 -0.0264 -0.0016 -0.0098 -0.0043 -0.0098 -0.0297 -0.0118 -0.0394 -0.0098 -0.0376 -0.0314 h g i H 0.0088 0.0088 0.0523 0.0122 0.0070 0.0078 0.0050 0.1586 0.0135 0.0135 0.0122 0.0679 0.0224 0.0109 0.0813 0.0117 -0.0179 -0.0179 -0.0363 -0.0179 -0.0285 -0.0029 -0.0125 -0.0285 Plateau Plateau -0.0385 -0.0385 -0.0127 -0.0127 -0.0385 -0.0225 -0.0065 -0.0407

Upper 0.2200 0.2200 0.0192 0.0014 0.0251 0.0251 0.0300 0.0463 0.0140 0.0815 -0.0308 -0.0308 -0.0377 -0.0622 -0.0217 -0.0086 -0.0308 -0.0453 -0.0357 -0.0173 -0.0234 -0.0252 -0.0149 -0.0149 -0.0173 -0.0217 -0.0375 -0.0217 -0.0059 -0.0098 -0.0594 -0.0634 Mesa Top Top Mesa

Lower 0.1063 0.1063 0.0232 0.0539 0.0539 0.0355 0.0101 0.0356 0.0252 -0.0352 -0.0352 -0.0431 -0.0711 -0.0248 -0.0007 -0.0352 -0.0514 -0.0236 -0.0400 -0.0240 -0.0192 -0.0147 -0.0054 -0.0054 -0.0248 -0.0372 -0.0248 -0.0051 -0.0248 -0.0755 -0.0184 -0.0797 Mesa Top Top Mesa 0.0012 0.0012 0.0629 0.0036 0.1977 0.0217 0.0119 0.0009 0.0009 0.0065 -0.0231 -0.0231 -0.0226 -0.0368 -0.0278 -0.0163 -0.0316 -0.0281 -0.0099 -0.0372 -0.0372 -0.0091 -0.0163 -0.0099 -0.0144 -0.0163 -0.0163 -0.0475 -0.0305 -0.0543 -0.0163 -0.0447 d an l Sands Slickrock Slickrock Sands 0.1887 0.1887 0.0153 0.0125 0.0125 0.0117 0.0727 Up -0.0295 -0.0295 -0.0335 -0.0432 -0.0108 -0.0240 -0.0192 -0.0030 -0.0540 -0.0051 -0.0131 -0.0251 -0.0175 -0.0368 -0.0368 -0.0389 -0.0240 -0.0177 -0.0507 -0.0240 -0.0240 -0.0709 -0.0238 -0.0175 -0.0708

y k Roc Slopes Slopes 0.0013 0.0013 0.0451 0.0673 0.0115 0.0115 0.0175 0.0152 0.0011 0.0676 -0.0303 -0.0303 -0.0469 -0.0188 -0.0368 -0.0114 -0.0342 -0.0295 -0.0107 -0.0105 -0.0153 -0.0621 -0.0005 -0.0005 -0.0368 -0.0220 -0.0609 -0.0287 -0.0290 -0.0609 -0.0177 -0.0649 id id Ar 0.0251 0.0251 0.2654 0.2274 0.0351 0.0370 0.0465 0.0465 0.0218 0.0220 0.2167 0.0839 0.0716 -0.0139 -0.0139 -0.0466 -0.0204 -0.0224 -0.0189 -0.0350 -0.0516 -0.0055 -0.0421 -0.0144 -0.0144 -0.0204 -0.0204 -0.0449 -0.0084 -0.0222 -0.0161 -0.0059 Lowlands Lowlands d an l Flats Flats 0.0099 0.0099 0.0310 0.0220 0.0436 0.0355 0.0355 0.0207 0.0290 0.0317 0.0388 0.2722 -0.0189 -0.0189 -0.0169 -0.0109 -0.0154 -0.0154 -0.0244 -0.0066 -0.0350 -0.0245 -0.0154 -0.0269 -0.0269 -0.0269 -0.0109 -0.0109 -0.0109 -0.0026 -0.0109 -0.0192 Low 3 8 8 1 2 2 5 4 5 2 5 9 6 1 1 6 1 1 1 9 6 1 19 10 27 22 10 Compositae Compositae Compositae Compositae Umbelliferae Betulaceae Gramineae Gramineae Gramineae Compositae Compositae Compositae Compositae Gramineae Chenopodiaceae 2 Asclepiadaceae Asclepiadaceae Compositae Compositae Compositae Compositae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Chenopodiaceae

3. Continued. sp.

sp. PPENDIX A Scientific Name Name Scientific Family # Occ. Atriplex confertifolia confertifolia Atriplex salicina Baccharis multiradiata Baileya sagittata Balsamorhiza erecta Berula Betula occidentalis curtipendula Bouteloua Bouteloua eriopoda Bouteloua gracilis Brickellia californica Brickellia grandiflora microphylla Brickellia Brickellia oblongifolia Bromus anomalus Asclepias subverticillata subverticillata Asclepias glaucodes Aster Aster Aster spathulatus welshii Aster Astragalus flavus Astragalus lancearius mollissimus Astragalus Astragalus nuttallianus Astragalus piutensis Astragalus subcinereus Astragalus Astragalus zionis canescens Atriplex 2015] PLANT COALITION GROUPS OF ZION NP 59

bank bank 0.0235 0.0235 0.2903 0.0094 0.2511 0.2511 0.0995 0.3353 0.0262 0.3576 0.0050 0.0067 0.0870 0.0708 0.0556 -0.0172 -0.0172 -0.0544 -0.0544 -0.0085 -0.0085 -0.0160 -0.0153 -0.0085 -0.0121 -0.0085 -0.0554 -0.0101 -0.0458 -0.0085 -0.0085 Stream-

ce on on i y 0.0264 0.0264 0.0118 0.0118 0.2602 0.0071 0.0179 0.0371 0.0547 0.0491 0.0179 0.0465 0.0042 0.0800 0.0260 0.0120 0.0430 Crev Can -0.0055 -0.0055 -0.0139 -0.0098 -0.0642 -0.0642 -0.0873 -0.0016 -0.0261 -0.0329 -0.0098 -0.0100 -0.0098 -0.0098 h g i H 0.0641 0.0641 0.0097 0.2714 0.2714 0.0473 0.0953 0.0074 0.0263 0.0403 0.0176 0.1027 0.0086 0.0230 0.0036 Plateau Plateau -0.0179 -0.0179 -0.0032 -0.0126 -0.0126 -0.0865 -0.1083 -0.0127 -0.0302 -0.0127 -0.0112 -0.0127 -0.0127 -0.0045 -0.0449 -0.0127

p

Upper 0.0788 0.0788 0.0053 0.0053 0.0060 0.1553 0.1229 0.0039 -0.0252 -0.0252 -0.0308 -0.0138 -0.0107 -0.0107 -0.0460 -0.0647 -0.1239 -0.1512 -0.0217 -0.0550 -0.0077 -0.0217 -0.0217 -0.0217 -0.0138 -0.0217 -0.0217 -0.0291 -0.0196 -0.0609 -0.0217 Mesa To Mesa

p

Lower 0.1121 0.1121 0.0107 0.0107 0.0014 0.2114 0.0218 0.0103 -0.0282 -0.0282 -0.0352 -0.0150 -0.0416 -0.0416 -0.0176 -0.0734 -0.1439 -0.1832 -0.0033 -0.0248 -0.0626 -0.0119 -0.0248 -0.0248 -0.0248 -0.0248 -0.0248 -0.0238 -0.0212 -0.0401 -0.0248 Mesa To Mesa 0.0757 0.0757 0.0454 0.0029 0.1222 0.0308 0.2903 -0.0174 -0.0174 -0.0196 -0.0196 -0.0056 -0.0372 -0.0372 -0.0407 -0.0855 -0.1214 -0.0099 -0.0163 -0.0395 -0.0340 -0.0100 -0.0006 -0.0113 -0.0099 -0.0163 -0.0022 -0.0003 -0.0163 -0.0163 d an l Sands Slickrock Slickrock Sands 0.0553 0.0553 0.0436 0.0098 0.0505 0.0315 0.0174 0.0831 Up -0.0180 -0.0180 -0.0143 -0.0290 -0.0094 -0.0456 -0.0456 -0.0486 -0.0613 -0.0990 -0.0590 -0.0149 -0.0240 -0.0562 -0.0240 -0.0169 -0.0169 -0.0240 -0.0240 -0.0227 -0.0240 -0.0240

y es es k p Roc Slo 0.0003 0.0003 0.0044 0.0731 0.0731 0.0710 0.0399 0.0000 0.0322 -0.0411 -0.0411 -0.0425 -0.0574 -0.0574 -0.0665 -0.0857 -0.0368 -0.0368 -0.0365 -0.0232 -0.0368 -0.0368 -0.0368 -0.0213 -0.0482 -0.0301 -0.0198 -0.0071 -0.0185 -0.0368 -0.0030 id id Ar 0.0051 0.0051 0.3680 0.3680 0.2135 0.1303 0.0121 0.0127 0.0040 -0.0389 -0.0389 -0.0197 -0.0198 -0.0434 -0.0434 -0.0670 -0.0336 -0.0162 -0.0204 -0.0117 -0.0204 -0.0204 -0.0204 -0.0204 -0.1431 -0.0347 -0.0260 -0.0182 -0.0544 -0.0117 -0.0154 Lowlands Lowlands d an l Flats Flats 0.0182 0.0182 0.0233 0.0017 0.0196 0.0293 0.0293 0.0312 0.0339 0.1156 0.0606 0.0171 0.0711 0.0606 0.0512 0.1112 -0.0196 -0.0196 -0.0109 -0.0109 -0.0154 -0.0109 -0.0109 -0.0109 -0.0109 -0.0335 -0.0291 -0.0154 -0.0392 -0.0109 Low 2 2 1 9 1 1 7 1 2 1 1 1 1 1 7 2 7 1 1 12 10 43 23 67 40 130 130 # Occ. Occ. # y Ulmaceae Ulmaceae Gramineae Gentianaceae Rhamnaceae 4 Rhamnaceae Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae Liliaceae Liliaceae Onagraceae Onagraceae Onagraceae Cruciferae Cyperaceae Cyperaceae Cyperaceae Cyperaceae Cyperaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Cruciferae Cruciferae

3. Continued. sp.

sp. PPENDIX A Scientific Name Name Scientific Famil Ceanothus martinii reticulata Celtis longispinus Cenchrus Centaurium exaltatum Bromus carinatus Bromus Bromus ciliatus Bromus diandrus* Bromus rubens* Bromus tectorum* Bromus vulgaris scopulorum Calamagrostis Calochortus flexuosus Calochortus nuttallii multijuga Camissonia parvula Camissonia Camissonia cordifolia Cardamine aurea Carex hystericina Carex occidentalis Carex rossii Carex Carex chromosa Castilleja Castilleja linariifolia scabrida Castilleja cooperi Caulanthus crassicaulis Caulanthus 60 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.0605 0.0605 0.0404 0.1104 0.0114 0.0487 0.0411 0.0911 0.0117 0.0120 -0.0149 -0.0149 -0.0535 -0.0521 -0.0586 -0.0085 -0.0085 -0.0065 -0.0149 -0.0085 -0.0192 -0.0085 -0.0085 -0.0211 -0.0196 -0.0315 -0.0121 -0.0095 -0.0505 -0.0085 S

ce i 0.0241 0.0241 0.1943 0.2611 0.0002 0.0364 0.0954 0.0035 0.0859 0.0337 Crev -0.0170 -0.0170 -0.0468 -0.0409 -0.0098 -0.0098 -0.0427 -0.0170 -0.0100 -0.0098 -0.0145 -0.0098 -0.0435 -0.0098 -0.0206 -0.0360 -0.0139 -0.0139 -0.0470 -0.0098 Canyon Canyon h g i H 0.1983 0.1983 0.0322 0.0581 0.0621 0.0337 0.0262 0.0514 0.0509 0.0123 0.0067 0.0544 -0.0220 -0.0220 -0.0540 -0.0299 -0.0622 -0.0127 -0.0065 -0.0518 -0.0220 -0.0127 -0.0127 -0.0592 -0.0127 -0.0711 -0.0032 -0.0466 -0.0146 -0.0127 Plateau Plateau

Upper 0.1440 0.1440 0.0032 0.0330 0.0045 0.0029 0.1618 0.0279 0.0182 0.0741 0.0349 0.0821 -0.0263 -0.0263 -0.0425 -0.0217 -0.0240 -0.0139 -0.0138 -0.0138 -0.0217 -0.0770 -0.0037 -0.0726 -0.0078 -0.0085 -0.0800 -0.0005 -0.0006 -0.0217 Mesa Top Top Mesa

Lower 0.2588 0.2588 0.1087 0.0486 0.0085 0.0140 0.0345 0.0987 0.0095 0.0028 0.0147 0.0343 0.0250 0.0257 -0.0431 -0.0431 -0.0254 -0.0008 -0.0248 -0.0075 -0.0203 -0.0150 -0.0248 -0.0952 -0.0023 -0.0941 -0.0914 -0.0060 -0.0142 -0.0248 Mesa Top Top Mesa 0.3022 0.3022 0.1140 0.0634 0.0370 0.0507 0.1966 0.1052 0.0046 0.0755 0.0116 0.0282 -0.0221 -0.0221 -0.0614 -0.0163 -0.0163 -0.0683 -0.0212 -0.0284 -0.0163 -0.0127 -0.0104 -0.0282 -0.0109 -0.0334 -0.0135 -0.0601 -0.0231 -0.0163 d an l Sands Slickrock Slickrock Sands 0.0948 0.0948 0.2527 0.0042 0.1933 0.1202 0.0130 0.0321 0.0434 0.0254 0.0087 0.1314 Up -0.0303 -0.0303 -0.0766 -0.0177 -0.0177 -0.0601 -0.0259 -0.0310 -0.0027 -0.0240 -0.0054 -0.0197 -0.0112 -0.0337 -0.0252 -0.0830 -0.0276 -0.0240

y k Roc Slopes Slopes 0.0125 0.0125 0.1114 0.0011 0.0804 0.0206 0.0285 0.0303 0.0050 0.0075 0.0063 0.0777 -0.0116 -0.0116 -0.0008 -0.0212 -0.0514 -0.0414 -0.0491 -0.0291 -0.0215 -0.0290 -0.0271 -0.0142 -0.0508 -0.0552 -0.0393 -0.0199 -0.0082 -0.0212 id id Ar 0.1220 0.1220 0.0466 0.0081 0.0881 0.0037 0.0045 0.0136 0.0138 0.2448 0.0019 0.0235 -0.0732 -0.0732 -0.0845 -0.0719 -0.0115 -0.0138 -0.0460 -0.0204 -0.0204 -0.0529 -0.0360 -0.0204 -0.0538 -0.0308 -0.0355 -0.0255 -0.0178 -0.0696 Lowlands Lowlands d an l Flats Flats 0.2085 0.2085 0.1195 0.0294 0.0518 0.0236 0.2224 0.0145 0.0286 -0.0189 -0.0189 -0.0741 -0.0411 -0.0625 -0.0109 -0.0109 -0.0189 -0.0189 -0.0245 -0.0109 -0.0109 -0.0433 -0.0109 -0.0081 -0.0109 -0.0006 -0.0154 -0.0219 -0.0686 -0.0109 Low 3 1 1 3 3 5 1 1 1 5 1 1 6 8 3 2 4 2 1 40 33 44 23 74 39 16 13 35 Polygonaceae Rosaceae Rosaceae Compositae Compositae Compositae Euphorbiaceae Euphorbiaceae Chenopodiaceae Pyrolaceae Pyrolaceae Cruciferae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Ranunculaceae Rosaceae Scrophulariaceae Polemoniaceae Polemoniaceae Santalaceae Convolvulaceae 3. Continued. PPENDIX A Centrostegia thurberi intricatusCercocarpus Cercocarpus montanus Chaenactis douglasii stevioides Chaenactis ericoides Chaetopappa albomarginata Chamaesyce fendleri Chamaesyce fremontii Chenopodium menziesii Chimaphila Chimaphila umbellata tenella* Chorispora jonesii Chrysopsis villosa Chrysopsis depressus Chrysothamnus greenei Chrysothamnus nauseosus Chrysothamnus Chrysothamnus parryi viscidiflorus Chrysothamnus arizonicum Cirsium wheeleri Cirsium columbiana Clematis ramosissima Coleogyne parviflora Collinsia Collomia grandiflora linearis Collomia umbellata Comandra arvensis*Convolvulus Scientific Name Name Scientific Family # Occ. 2015] PLANT COALITION GROUPS OF ZION NP 61

bank bank tream- 0.0050 0.0050 0.0264 0.0264 0.0691 0.0686 0.2023 0.0103 0.0494 0.1908 S -0.0342 -0.0342 -0.0085 -0.0149 -0.0085 -0.0121 -0.0121 -0.0325 -0.0149 -0.0469 -0.0121 -0.0085 -0.0639 -0.0085 -0.0085 -0.0339 -0.0085 -0.0315 -0.0085 -0.0141 -0.0085 -0.0214

ce i 0.0963 0.0963 Crev 0.0260 0.0260 0.3184 0.2342 0.0131 Canyon Canyon -0.0440 -0.0440 -0.0098 -0.0170 -0.0098 -0.0139 -0.0139 -0.0398 -0.0170 -0.0403 -0.0139 -0.0098 -0.0776 -0.0098 -0.0098 -0.0388 -0.0098 -0.0020 -0.0360 -0.0073 -0.0098 -0.0172 -0.0098 -0.0520 -0.0016 h g i H 0.0096 0.0096 0.0406 0.0406 0.0007 0.0764 0.3368 0.0007 0.0040 0.0264 0.0116 0.0787 0.0632 Plateau Plateau -0.0393 -0.0393 -0.0127 -0.0178 -0.0127 -0.0129 -0.0129 -0.0226 -0.0178 -0.0683 -0.0131 -0.0127 -0.0746 -0.0127 -0.0351 -0.0127 -0.0412 -0.0138 -0.0127

Upper 0.0030 0.0030 0.0171 0.0171 0.0713 0.0009 0.0510 0.0500 0.0115 -0.0138 -0.0138 -0.0124 -0.0003 -0.0059 -0.0112 -0.0112 -0.0341 -0.0204 -0.0637 -0.0252 -0.0028 -0.0796 -0.0217 -0.0119 -0.0217 -0.0366 -0.0570 -0.0332 -0.0119 -0.0588 -0.0081 -0.0217 Mesa Top Top Mesa

Lower 0.0145 0.0145 0.0103 0.1293 0.1293 0.0741 0.0076 0.0404 -0.0150 -0.0150 -0.0132 -0.0193 -0.0212 -0.0212 -0.0166 -0.0466 -0.0301 -0.0691 -0.0282 -0.0248 -0.1192 -0.0248 -0.0248 -0.0248 -0.0160 -0.0129 -0.0649 -0.0374 -0.0248 -0.0556 -0.0150 -0.0248 Mesa Top Top Mesa 0.0018 0.0018 0.0972 0.0972 0.0833 0.0665 -0.0655 -0.0655 -0.0163 -0.0054 -0.0104 -0.0092 -0.0092 -0.0065 -0.0157 -0.0223 -0.0231 -0.0163 -0.0596 -0.0163 -0.0083 -0.0408 -0.0163 -0.0231 -0.0565 -0.0284 -0.0083 -0.0293 -0.0045 -0.0819 -0.0163 -0.0113 d an l Sands Slickrock Slickrock Sands Up 0.0580 0.0580 0.0443 0.0443 0.2622 0.0253 0.0887 0.0211 0.1042 0.1042 -0.0369 -0.0369 -0.0240 -0.0240 -0.0097 -0.0101 -0.0101 -0.0251 -0.0240 -0.0240 -0.0356 -0.0240 -0.0311 -0.0211 -0.0688 -0.0417 -0.0119 -0.0007 -0.0494 -0.0165 -0.0337 -0.0169

y k Roc Slopes Slopes 0.0772 0.0772 0.0669 0.0150 0.0314 0.0358 0.0358 0.1252 0.1317 0.0414 0.0497 0.0237 0.0414 0.0713 -0.0368 -0.0368 -0.0085 -0.0085 -0.0006 -0.0137 -0.0284 -0.0095 -0.0368 -0.0212 -0.0547 -0.0466 -0.0595 -0.0169 -0.0142 -0.0279 -0.0482 -0.0287 id id Ar 0.0745 0.0745 0.0330 0.0151 0.0247 0.0247 0.0704 0.2085 0.0333 0.0499 0.1213 0.1716 0.0039 -0.0204 -0.0204 -0.0118 -0.0156 -0.0156 -0.0224 -0.0177 -0.0031 -0.0111 -0.0117 -0.0106 -0.0505 -0.0239 -0.0306 -0.0106 -0.0339 -0.0084 -0.0204 -0.0411 Lowlands Lowlands d an l Flats Flats 0.0411 0.0411 0.0018 0.0286 0.1155 0.0133 0.0155 0.0076 0.0779 -0.0072 -0.0072 -0.0109 -0.0109 -0.0189 -0.0109 -0.0154 -0.0154 -0.0462 -0.0351 -0.0109 -0.0026 -0.0109 -0.0240 -0.0154 -0.0049 -0.0109 -0.0061 -0.0037 -0.0109 -0.0219 -0.0109 Low 1 1 3 1 2 3 2 1 1 1 1 6 2 3 1 7 1 1 4 1 19 17 31 60 15 13 28 Scrophulariaceae Scrophulariaceae Scrophulariaceae Cornaceae Compositae Compositae Compositae Compositae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Boraginaceae Cucurbitaceae Umbelliferae Umbelliferae Umbelliferae Polypodiaceae Gramineae Leguminosae Solanaceae Ranunculaceae Ranunculaceae Cruciferae Cruciferae Cruciferae Liliaceae Gramineae Compositae 2 Compositae

sp. sp. 3. Continued. sp. (annual) sp.

sp. PPENDIX A Scientific Name Name Scientific Family # Occ. Cordylanthus kingii Cordylanthus Cordylanthus sericea Cornus acuminata Crepis Crepis occidentalis Crepis runcinata Crepis Crepis Cryptantha cinerea Cryptantha circumscissa Cryptantha confertiflora Cryptantha fendleri humilis Cryptantha Cryptantha foetidissima Cucurbita newberryi Cymopterus purpureus Cymopterus Cymopterus fragilis Cystopteris glomerata* Dactylis Dalea searlsiae wrightii Datura Delphinium andersonii Delphinium nuttallianum californica Descurainia pinnata Descurainia sophia* Descurainia trachycarpum Disporum Distichlis spicata 62 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.2409 0.2409 0.0390 0.0707 0.1360 0.0103 0.3576 0.3744 0.3213 0.0094 0.0094 0.0817 0.1572 0.1572 0.2742 0.0127 -0.0085 -0.0085 -0.0055 -0.0172 -0.0374 -0.0303 -0.0085 -0.0025 -0.0025 -0.0142 -0.0189 -0.0085 -0.0229 -0.0377 -0.0314 -0.0085 S

ce i 0.0698 0.0698 0.1830 0.1889 0.2225 0.0179 0.0371 0.1603 0.0132 0.0387 0.0387 0.1409 0.2531 0.0117 0.0117 0.2267 Crev Canyon Canyon -0.0098 -0.0098 -0.0167 -0.0197 -0.0394 -0.0366 -0.0016 -0.0098 -0.0098 -0.0615 -0.0098 -0.0261 -0.0394 -0.0247 -0.0043 -0.0170 h g i H 0.0236 0.0236 0.1829 0.0187 0.0497 0.0690 0.0787 0.0263 0.0372 0.0097 0.0148 0.0148 0.0479 0.0122 0.0122 0.0581 0.0583 Plateau Plateau -0.0091 -0.0091 -0.0255 -0.0554 -0.0466 -0.0127 -0.0025 -0.0025 -0.0115 -0.0032 -0.0737 -0.0302 -0.0488 -0.0206 -0.0127

Upper 0.0116 0.0116 0.0115 0.0067 0.0067 0.0309 -0.0099 -0.0099 -0.0244 -0.0436 -0.0854 -0.0326 -0.0286 -0.0286 -0.0286 -0.0217 -0.0217 -0.0537 -0.0308 -0.0086 -0.0086 -0.0138 -0.0745 -0.0040 -0.0186 -0.0520 -0.0232 -0.0217 -0.0217 -0.0357 -0.0274 -0.0059 Mesa Top Top Mesa

Lower 0.0027 0.0027 0.0018 0.0802 0.0802 0.0014 0.0006 -0.0100 -0.0100 -0.0503 -0.0499 -0.0888 -0.0459 -0.0317 -0.0055 -0.0150 -0.0248 -0.0248 -0.0501 -0.0352 -0.0123 -0.0123 -0.0150 -0.0816 -0.0588 -0.0478 -0.0248 -0.0248 -0.0254 -0.0536 -0.0051 -0.0119 Mesa Top Top Mesa 0.0420 0.0420 0.0404 0.2068 0.2068 0.0224 0.0065 0.0062 0.0141 -0.0111 -0.0111 -0.0193 -0.0399 -0.0328 -0.0630 -0.0456 -0.0058 -0.0163 -0.0163 -0.0113 -0.0190 -0.0231 -0.0108 -0.0108 -0.0056 -0.0704 -0.0113 -0.0267 -0.0163 -0.0163 -0.0053 -0.0115 d an l Sands Slickrock Slickrock Sands 0.0153 0.0153 0.0110 0.0971 0.0971 0.0050 0.0608 0.0133 Up -0.0046 -0.0046 -0.0271 -0.0351 -0.0451 -0.0872 -0.0253 -0.0060 -0.0165 -0.0240 -0.0169 -0.0464 -0.0340 -0.0340 -0.0340 -0.0094 -0.0141 -0.0189 -0.0347 -0.0240 -0.0240 -0.0170 -0.0026 -0.0185

y k Roc Slopes Slopes 0.0091 0.0091 0.0730 0.0124 0.0124 0.0044 0.0867 0.0162 0.1300 0.0099 0.0146 -0.0177 -0.0177 -0.0195 -0.0334 -0.0583 -0.0461 -0.0595 -0.0417 -0.0279 -0.0368 -0.0232 -0.0812 -0.0466 -0.0469 -0.0469 -0.0216 -0.0467 -0.0368 -0.0368 -0.0516 -0.0420 id id Ar 0.1980 0.1980 0.2637 0.0204 0.0051 0.0051 0.2595 0.0998 0.1003 0.0923 -0.0139 -0.0139 -0.0048 -0.0334 -0.0331 -0.0355 -0.0355 -0.0204 -0.0204 -0.0117 -0.0422 -0.0260 -0.0290 -0.0290 -0.0823 -0.0110 -0.0290 -0.0204 -0.0204 -0.0346 -0.0116 -0.0112 Lowlands Lowlands d an l Flats Flats 0.0722 0.0722 0.0475 0.0478 0.0225 0.0242 0.0196 0.0196 0.0217 0.0207 0.0099 0.0162 0.1929 -0.0109 -0.0109 -0.0042 -0.0215 -0.0189 -0.0006 -0.0109 -0.0109 -0.0109 -0.0154 -0.0154 -0.0154 -0.0516 -0.0154 -0.0407 -0.0109 -0.0109 -0.0219 -0.0109 Low 6 4 3 3 3 1 1 1 8 2 2 1 1 2 7 1 1 4 1 3 10 18 16 33 40 22 58 Gramineae Gramineae Compositae Ephedraceae Ephedraceae Onagraceae Onagraceae Onagraceae Onagraceae Orchidaceae Equisetaceae Equisetaceae Equisetaceae Gramineae 2 Primulaceae Primulaceae Cruciferae Cruciferae Cruciferae Labiatae Polypodiaceae Cactaceae Cactaceae Cyperaceae Cyperaceae Gramineae Gramineae Gramineae Gramineae

3. Continued.

sp. sp. PPENDIX A Scientific Name Name Scientific Family # Occ. Elymus Elymus spicatus Elymus trachycaulus Elymus frutescens Encelia Ephedra nevadensis viridis Ephedra Epilobium brachycarpum Epilobium ciliatum Epilobium glandulosum Epilobium gigantea Epipactis arvense Equisetum hyemale Equisetum laevigatum Equisetum Dodecatheon pulchellum Dodecatheon Draba asprella Draba cuneifolia verna* Draba Dracocephalum parviflorum filix-mas Dryopteris engelmannii Echinocereus triglochidiatus Echinocereus Eleocharis palustris parishii Eleocharis Elymus canadensis elymoides Elymus hispidus* Elymus smithii Elymus 2015] PLANT COALITION GROUPS OF ZION NP 63

bank bank tream- 0.0462 0.0462 0.2472 0.2472 0.0426 0.0674 0.0143 -0.0121 -0.0121 -0.0211 -0.0527 -0.0192 -0.0229 -0.0229 -0.0250 -0.0316 -0.0170 -0.0085 -0.0085 -0.0560 -0.0211 -0.0085 -0.0478 -0.0149 -0.0085 -0.0149 -0.0275 -0.0290 -0.0339 -0.0253 -0.0229 -0.0264 S

ce i y 0.3612 0.3612 0.0259 0.0259 0.0885 0.1662 0.0046 0.0517 0.0074 Crev -0.0139 -0.0139 -0.0282 -0.0287 -0.0220 -0.0261 -0.0261 -0.0360 -0.0393 -0.0043 -0.0098 -0.0246 -0.0588 -0.0147 -0.0098 -0.0503 -0.0170 -0.0098 -0.0170 -0.0065 -0.0305 -0.0369 -0.0261 Canyon Canyon h g i H 0.0630 0.0630 0.0552 0.0552 0.0184 0.0007 0.0187 0.1538 0.0394 -0.0179 -0.0179 -0.0407 -0.0315 -0.0243 -0.0339 -0.0339 -0.0466 -0.0192 -0.0127 -0.0216 -0.0713 -0.0224 -0.0127 -0.0592 -0.0220 -0.0127 -0.0220 -0.0361 -0.0245 -0.0503 -0.0339 -0.0145 Plateau Plateau p

Upper 0.0369 0.0369 0.0060 0.0173 0.0173 0.0284 0.0560 0.0804 0.0302 0.0085 -0.0308 -0.0308 -0.0609 -0.0365 -0.0550 -0.0550 -0.0710 -0.0131 -0.0252 -0.0059 -0.0145 -0.0681 -0.0125 -0.0217 -0.0554 -0.0128 -0.0019 -0.0377 -0.0502 -0.0519 -0.0779 -0.0430 Mesa Top Top Mesa p

Lower 0.0100 0.0100 0.0054 0.0515 0.0168 0.0168 0.0233 0.0074 0.0253 0.0479 0.0043 -0.0352 -0.0352 -0.0766 -0.0626 -0.0626 -0.0914 -0.0115 -0.0077 -0.0051 -0.0700 -0.0118 -0.0248 -0.0761 -0.0024 -0.0431 -0.0553 -0.0589 -0.0797 -0.0038 -0.0475 -0.0200 Mesa Top Top Mesa 0.0332 0.0332 0.1492 0.2338 0.2338 0.0322 0.0029 0.2217 0.0216 0.2322 0.0704 0.0862 -0.0488 -0.0488 -0.0076 -0.0217 -0.0417 -0.0417 -0.0495 -0.0587 -0.0053 -0.0271 -0.0154 -0.0274 -0.0163 -0.0310 -0.0206 -0.0029 -0.0284 -0.0378 -0.0476 -0.0297 d an l Sands Slickrock Slickrock Sands 0.1601 0.1601 0.0127 0.1198 0.1198 0.0062 0.0211 0.0569 0.0464 0.2047 0.0399 0.0856 0.0466 0.1001 0.1447 Up -0.0208 -0.0208 -0.0748 -0.0391 -0.0395 -0.0395 -0.0568 -0.0308 -0.0103 -0.0026 -0.0328 -0.0224 -0.0180 -0.0381 -0.0408 -0.0719 -0.0002

y k p Roc Slopes Slopes 0.2064 0.2064 0.0095 0.0095 0.0203 0.0146 0.0122 0.2246 0.1227 0.0748 0.0645 0.0316 0.0990 0.0162 0.0243 -0.0356 -0.0356 -0.0910 -0.0670 -0.0518 -0.0212 -0.0212 -0.0209 -0.0127 -0.0138 -0.0358 -0.0374 -0.0297 -0.0251 -0.0216 -0.0459 -0.0469 id id Ar 0.1098 0.1098 0.1277 0.2058 0.2058 0.1752 0.1342 0.0237 0.1334 0.0211 0.1657 0.0652 0.2175 0.0333 -0.0340 -0.0340 -0.0709 -0.0382 -0.0719 -0.0719 -0.0041 -0.0198 -0.0116 -0.0001 -0.0387 -0.0196 -0.0409 -0.0291 -0.0204 -0.0338 -0.0898 -0.0305 Lowlands Lowlands d an l Flats Flats 0.0099 0.0099 0.2865 0.0414 0.0561 0.0561 0.0183 0.0224 0.0057 0.0610 0.0519 0.0092 -0.0775 -0.0775 -0.0081 -0.0062 -0.0062 -0.0448 -0.0097 -0.0154 -0.0109 -0.0109 -0.0219 -0.0341 -0.0269 -0.0379 -0.0189 -0.0109 -0.0350 -0.0291 -0.0291 -0.0382 Low 5 7 2 1 1 4 6 6 4 3 1 3 7 10 64 12 13 16 17 14 41 32 10 15 15 59 21 y Geraniaceae Cruciferae Compositae Euphorbiaceae Gramineae 2 Polemoniaceae Polemoniaceae Polemoniaceae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Compositae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae Polygonaceae

3. Continued.

sp. sp. PPENDIX A Erioneuron pilosum Erioneuron cicutarium* Erodium asperum Erysimum Eupatorium herbaceum brachycera Euphorbia Scientific Name Name Scientific Family # Occ. Eriastrum eremicum eremicum Eriastrum sparsiflorum Eriastrum canaani Erigeron divergens Erigeron flagellaris Erigeron pumilus Erigeron religious Erigeron Erigeron sionis Erigeron speciosus Erigeron utahensis Erigeron Eriogonum alatum corymbosum Eriogonum davidsonii Eriogonum fasciculatum Eriogonum heracleoides Eriogonum Eriogonum inflatum jamesii Eriogonum microthecum Eriogonum palmerianum Eriogonum racemosum Eriogonum Eriogonum Eriogonum umbellatum 64 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.1862 0.1862 0.0020 0.0290 0.0247 0.0264 0.0562 0.0207 0.1104 0.0114 0.0465 0.0404 0.1083 -0.0085 -0.0085 -0.0085 -0.0469 -0.0520 -0.0260 -0.0249 -0.0085 -0.0260 -0.0085 -0.0584 -0.0085 -0.0085 -0.0086 -0.0192 -0.0085 -0.0085 S

ce i 0.0263 0.0263 0.0227 0.0041 0.0520 0.0890 0.2616 0.2611 0.0002 0.1052 0.2131 -0.0098 -0.0098 -0.0030 -0.0819 -0.0092 -0.0153 -0.0068 -0.0098 -0.0226 -0.0098 -0.0272 -0.0969 -0.0098 -0.0098 -0.0962 -0.0125 -0.0098 -0.0081 -0.0098 Crev Canyon Canyon h g i H 0.0417 0.0417 0.2181 0.0003 0.0434 0.1380 0.0581 0.0014 0.0031 0.0882 0.0411 -0.0127 -0.0127 -0.0127 -0.0946 -0.0571 -0.0130 -0.0235 -0.0179 -0.0183 -0.0065 -0.0232 -0.0127 -0.1140 -0.0127 -0.0127 -0.1043 -0.0127 -0.0255 -0.0127 Plateau Plateau

Upper 0.0135 0.0135 0.0020 0.0025 0.0362 0.0777 0.0257 -0.0020 -0.0020 -0.1123 -0.0496 -0.0252 -0.0191 -0.0059 -0.0040 -0.0252 -0.0310 -0.0135 -0.0217 -0.0138 -0.0963 -0.0059 -0.0138 -0.0305 -0.0076 -0.0217 -0.1114 -0.0057 -0.0397 -0.0138 Mesa Top Top Mesa

Lower 0.0473 0.0473 0.0824 0.0121 0.0014 0.0039 0.0300 0.0238 0.0140 0.0112 0.0369 0.1376 -0.0248 -0.0248 -0.1467 -0.0546 -0.0199 -0.0282 -0.0195 -0.0130 -0.0196 -0.0248 -0.1116 -0.0150 -0.0346 -0.0248 -0.1590 -0.0110 -0.0449 -0.0150 Mesa Top Top Mesa 0.0168 0.0168 0.0183 0.0953 0.0406 0.0169 0.0370 0.0870 0.0260 -0.0163 -0.0163 -0.0069 -0.0313 -0.0179 -0.0231 -0.0202 -0.0185 -0.0163 -0.0256 -0.0163 -0.0406 -0.1029 -0.0163 -0.0163 -0.0163 -0.0984 -0.0109 -0.0163 -0.0278 -0.0106 d an l Sands Slickrock Slickrock Sands 0.0993 0.0993 0.1442 0.0320 0.0515 0.0269 0.0485 0.0033 Up -0.0240 -0.0240 -0.0240 -0.0204 -0.0478 -0.0340 -0.0275 -0.0185 -0.0020 -0.0006 -0.0375 -0.0030 -0.0240 -0.0369 -0.0027 -0.0240 -0.0374 -0.0240 -0.0696 -0.0174 -0.0411 -0.0240

y k Roc Slopes Slopes 0.0064 0.0064 0.0657 0.2408 0.0404 0.1091 0.0788 0.0132 0.0376 0.1513 0.0031 0.0182 0.0670 0.0180 0.0151 -0.0293 -0.0293 -0.0638 -0.0413 -0.0198 -0.0306 -0.0340 -0.0287 -0.0370 -0.0291 -0.0215 -0.0690 -0.0368 -0.0294 -0.0699 id id Ar 0.0035 0.0035 0.2480 0.0431 0.0830 0.0007 0.1296 0.2558 0.0004 0.0037 0.2817 0.0772 -0.0204 -0.0204 -0.0349 -0.0276 -0.0180 -0.0290 -0.0715 -0.0194 -0.0174 -0.0363 -0.0316 -0.0204 -0.0003 -0.0460 -0.0117 -0.0146 -0.0204 -0.0202 Lowlands Lowlands d an l Flats Flats 0.3561 0.3561 0.0169 0.0770 0.0411 0.0196 0.0249 0.0125 0.1195 0.0449 0.0989 0.0219 0.1243 0.0207 -0.0109 -0.0109 -0.0588 -0.0169 -0.0225 -0.0144 -0.0557 -0.0154 -0.0219 -0.0331 -0.0109 -0.0137 -0.0109 -0.0331 -0.0245 -0.0109 Low 1 1 6 9 2 6 2 1 2 4 9 1 1 1 1 9 5 1 5 1 4 1 74 45 28 14 84 94 Euphorbiaceae Euphorbiaceae Saxifragaceae Gramineae Oleaceae Oleaceae Liliaceae Compositae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Rubiaceae Garryaceae Onagraceae Onagraceae Geraniaceae Polemoniaceae Polemoniaceae Leguminosae Compositae Orchidaceae Compositae Compositae Boraginaceae Compositae Compositae Compositae

sp. 3. Continued.

sp. PPENDIX A Euphorbia spathulata utahensis Fendlerella Festuca octoflora Fraxinus anomala Fraxinus velutina Fritillaria atropurpurea Gaillardia pinnatifida Galium aparine Galium bifolium multiflorum Galium Galium Galium trifidum Galium triflorum flavescens Garrya coccinea Gaura Gayophytum richardsonii Geranium Gilia inconspicua Gilia scopulorum Glycyrrhiza lepidota Gnaphalium wrightii oblongifolia Goodyera Grindelia squarrosa spp. Gutierrezia patens Hackelia armerioides Haplopappus Haplopappus gracilis laricifolius Haplopappus Scientific Name Name Scientific Family # Occ. 2015] PLANT COALITION GROUPS OF ZION NP 65

bank bank tream- 0.0057 0.0057 0.0166 0.0538 0.3103 0.2580 0.3576 0.0109 0.0185 0.0721 -0.0515 -0.0515 -0.0085 -0.0121 -0.0121 -0.0007 -0.0085 -0.0510 -0.0222 -0.0085 -0.0085 -0.0025 -0.0242 -0.0192 -0.0112 -0.0024 -0.0229 -0.1076 -0.0085 -0.0121 S

ce i 0.2533 0.2533 0.0131 0.1947 0.1573 0.0105 0.1444 0.0552 0.0611 0.0371 0.0134 0.0243 0.1780 -0.0055 -0.0055 -0.0098 -0.0139 -0.0139 -0.0139 -0.0675 -0.0098 -0.0098 -0.0309 -0.0183 -0.0220 -0.0070 -0.0098 -0.1273 -0.0098 -0.0028 Crev Canyon Canyon h g i H 0.0476 0.0476 0.0432 0.0033 0.0007 0.0325 0.0472 0.0263 0.1754 0.0344 0.0166 0.0516 0.0628 -0.0509 -0.0509 -0.0058 -0.0179 -0.0179 -0.0021 -0.0055 -0.0683 -0.0027 -0.0127 -0.0065 -0.0122 -0.0049 -0.0114 -0.0127 -0.0284 -0.1258 Plateau Plateau

Upper 0.0324 0.0324 0.1300 0.1539 0.1344 0.0318 0.0409 0.0359 0.0009 -0.0308 -0.0308 -0.0168 -0.0019 -0.0045 -0.1138 -0.0308 -0.0020 -0.0119 -0.0130 -0.0248 -0.0196 -0.0247 -0.0217 -0.0318 -0.0252 -0.0377 -0.0252 -0.0217 -0.0387 -0.0126 Mesa Top Top Mesa

Lower 0.0255 0.0255 0.0461 0.0860 0.0382 0.0093 0.0165 0.0541 0.0226 0.0539 0.0235 0.0109 -0.0352 -0.0352 -0.0352 -0.1262 -0.0046 -0.0248 -0.0248 -0.0134 -0.0291 -0.0169 -0.0248 -0.0287 -0.0282 -0.0431 -0.0166 -0.0248 -0.0521 -0.0129 Mesa Top Top Mesa 0.0046 0.0046 0.1744 0.0327 0.3078 0.0072 0.0522 0.0315 0.2693 0.0206 0.0182 -0.0099 -0.0099 -0.0231 -0.0231 -0.0185 -0.0871 -0.0163 -0.0077 -0.0263 -0.0292 -0.0163 -0.0143 -0.0150 -0.0113 -0.1486 -0.0283 -0.0109 -0.0233 -0.0116 d an l Sands Slickrock Slickrock Sands 0.0711 0.0711 0.0744 0.0077 0.0480 0.2352 0.0043 0.1277 0.0594 0.0052 0.0017 Up -0.0165 -0.0165 -0.0340 -0.0340 -0.0193 -0.0292 -0.1195 -0.0240 -0.0251 -0.0390 -0.0240 -0.0290 -0.0180 -0.0290 -0.0169 -0.0462 -0.0170 -0.0662 -0.0223

y k Roc Slopes Slopes 0.1804 0.1804 0.0402 0.0185 0.0435 0.0396 0.1868 -0.0052 -0.0052 -0.0522 -0.0468 -0.0069 -0.0461 -0.0138 -0.0438 -0.0293 -0.0049 -0.0359 -0.0048 -0.0368 -0.0373 -0.0466 -0.0471 -0.0425 -0.0232 -0.0461 -0.0217 -0.0836 -0.0359 -0.0326 id id Ar 0.2507 0.2507 0.0035 0.0044 0.0776 0.0176 0.1706 -0.0438 -0.0438 -0.0161 -0.0166 -0.0058 -0.0290 -0.0709 -0.0156 -0.0921 -0.0290 -0.0381 -0.0395 -0.0117 -0.0435 -0.0260 -0.0305 -0.0228 -0.0117 -0.0775 -0.0159 -0.0045 -0.0223 -0.0143 Lowlands Lowlands d an l Flats Flats 0.0241 0.0241 0.3310 0.1280 0.3561 0.1070 0.0352 0.0251 0.0689 0.2237 0.0069 0.0352 -0.0620 -0.0620 -0.0109 -0.0154 -0.0292 -0.0109 -0.0356 -0.0109 -0.0154 -0.0368 -0.0027 -0.0417 -0.0109 -0.0291 -0.0189 -0.0154 -0.0109 -0.0178 Low 1 2 2 1 1 1 2 5 5 1 7 2 3 2 1 1 2 2 42 12 41 21 11 14 21 13 115 115 Compositae Compositae Compositae Compositae Compositae Saxifragaceae Compositae Gramineae Rosaceae Gramineae Gramineae Hydrophyllaceae Compositae Compositae Polemoniaceae Polemoniaceae Compositae Rosaceae Saxifragaceae Juncaceae Juncaceae Juncaceae Cupressaceae Cupressaceae Gramineae Compositae Compositae Boraginaceae Compositae 2 Compositae occidentale

3. Continued.

sp. sp. PPENDIX A Haplopappus scopulorum Helianthella quinquenervis Helianthus annuus Helianthus anomalus Helianthus nuttallii rubescens Heuchera fendleri Hieracium Hilaria dumosus Holodiscus murinum* Hordeum Hordeum Hydrophyllum filifolius Hymenopappus cooperi Hymenoxys aggregata Ipomopsis congesta Ipomopsis Iva axillaris sabulosa Ivesia americana Jamesia Juncus arcticus Juncus ensifolius Juncus tenuis osteosperma Juniperus scopulorum Juniperus macrantha Koeleria Lactuca serriola* tatarica Lactuca Lappula occidentalis Scientific Name Name Scientific Family # Occ. 66 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.0067 0.0067 0.0006 0.0000 0.0352 0.0067 0.0067 0.0114 0.0390 -0.0134 -0.0134 -0.0121 -0.0085 -0.0085 -0.0211 -0.0363 -0.0162 -0.0085 -0.0172 -0.0085 -0.0121 -0.0260 -0.0260 -0.0321 -0.0121 -0.0245 -0.0085 -0.0196 -0.0275 -0.0085 -0.0090 -0.0265 S

ce i 0.0071 0.0071 0.1068 0.0136 0.0136 0.0002 0.0551 0.0196 0.0617 Crev -0.0012 -0.0012 -0.0139 -0.0080 -0.0067 -0.0098 -0.0242 -0.0374 -0.0305 -0.0098 -0.0197 -0.0098 -0.0139 -0.0064 -0.0297 -0.0297 -0.0253 -0.0100 -0.0254 -0.0098 -0.0098 -0.0098 -0.0335 Canyon Canyon h g i H 0.0257 0.0257 0.0680 0.1080 0.0413 0.0350 0.2937 0.0463 0.0463 0.1108 0.0870 -0.0255 -0.0255 -0.0131 -0.0127 -0.0285 -0.0514 -0.0127 -0.0255 -0.0127 -0.0036 -0.0323 -0.0323 -0.0112 -0.0341 -0.0127 -0.0127 -0.0213 -0.0149 -0.0127 -0.0092 -0.0485 Plateau Plateau

Upper 0.0333 0.0333 0.0026 0.0386 0.0358 0.1321 0.0474 0.0436 0.2432 0.2432 0.0053 0.0053 0.0036 0.0042 -0.0059 -0.0059 -0.0045 -0.0138 -0.0235 -0.0524 -0.0217 -0.0298 -0.0217 -0.0064 -0.0064 -0.0124 -0.0466 -0.0217 -0.0138 -0.0311 -0.0309 -0.0666 Mesa Top Top Mesa

Lower 0.0372 0.0372 0.0576 0.0256 0.0389 0.0016 0.0058 0.0058 0.1446 0.0105 0.0076 -0.0051 -0.0051 -0.0027 -0.0035 -0.0150 -0.0439 -0.0600 -0.0248 -0.0449 -0.0248 -0.0002 -0.0122 -0.0122 -0.0033 -0.0559 -0.0248 -0.0150 -0.0033 -0.0228 -0.0110 -0.0784 Mesa Top Top Mesa 0.0469 0.0469 0.0469 0.0010 0.0010 0.2160 0.0732 0.1762 -0.0163 -0.0163 -0.0231 -0.0150 -0.0269 -0.0035 -0.0379 -0.0188 -0.0268 -0.0056 -0.0303 -0.0056 -0.0231 -0.0299 -0.0346 -0.0346 -0.0433 -0.0090 -0.0163 -0.0344 -0.0163 -0.0163 -0.0045 -0.0175 d an l Sands Slickrock Slickrock Sands 0.0251 0.0251 0.0115 0.0034 0.0393 0.0133 0.0161 0.0161 0.0261 0.0080 0.0210 0.1760 Up -0.0149 -0.0149 -0.0246 -0.0296 -0.0240 -0.0513 -0.0260 -0.0106 -0.0376 -0.0142 -0.0294 -0.0259 -0.0259 -0.0158 -0.0240 -0.0499 -0.0177 -0.0240 -0.0073 -0.0073

y k Roc Slopes Slopes 0.0699 0.0699 0.0067 0.1080 0.0057 0.0141 0.0917 0.0917 0.0088 0.0209 0.0119 0.0315 0.0400 -0.0067 -0.0067 -0.0033 -0.0354 -0.0290 -0.0063 -0.0177 -0.0165 -0.0162 -0.0176 -0.0838 -0.0283 -0.0283 -0.0294 -0.0475 -0.0107 -0.0215 -0.0290 -0.0113 id id Ar 0.0099 0.0099 0.0262 0.1402 0.0903 0.0659 0.0722 0.0136 0.0319 0.0319 0.1408 0.0550 0.0037 0.0074 0.0073 0.1643 -0.0198 -0.0198 -0.0071 -0.0114 -0.0389 -0.0204 -0.0159 -0.0308 -0.0536 -0.0063 -0.0063 -0.0783 -0.0204 -0.0144 -0.0489 -0.0681 Lowlands Lowlands d an l Flats Flats 0.0017 0.0017 0.0524 0.1489 0.0040 0.0692 0.1484 0.1484 0.0196 0.1195 0.0537 0.0000 -0.0070 -0.0070 -0.0109 -0.0219 -0.0109 -0.0109 -0.0171 -0.0109 -0.0219 -0.0109 -0.0154 -0.0331 -0.0331 -0.0150 -0.0154 -0.0109 -0.0312 -0.0350 -0.0109 -0.0433 Low 1 2 2 4 1 1 6 1 4 1 2 9 9 2 1 8 1 1 1 8 1 17 12 20 10 19 14 Saxifragaceae Saxifragaceae Boraginaceae Boraginaceae Umbelliferae Umbelliferae Umbelliferae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Leguminosae Linaceae 6 Linaceae Leguminosae Leguminosae Leguminosae Compositae Cruciferae Cruciferae Cruciferae Cruciferae Cruciferae Polemoniaceae Polemoniaceae Liliaceae Polemoniaceae Polemoniaceae

3. Continued. sp. PPENDIX A Linum perenne* perenne* Linum Lithophragma tenellum incisum Lithospermum ruderale Lithospermum dissectum Lomatium graveolens Lomatium Lomatium Lotus denticulatus Lotus plebeius Lotus utahensis Lupinus argenteus brevicaulis Lupinus concinnus Lupinus kingii Lupinus sericeus Lupinus Scientific Name Name Scientific Family # Occ. Lathyrus brachycalyx Lathyrus lanszwertii Lathyrus Layia glandulosa campestre* Lepidium densiflorum Lepidium fremontii Lepidium lasiocarpum Lepidium montanum Lepidium pungens Leptodactylon watsonii Leptodactylon montanum Leucocrinum Linanthastrum nuttallii dichotomus Linanthus 2015] PLANT COALITION GROUPS OF ZION NP 67

bank bank tream- 0.2180 0.2180 0.0806 0.0067 0.0067 0.0000 0.2964 0.0723 0.1908 0.4632 0.1045 0.0442 0.1089 0.0167 -0.0121 -0.0121 -0.0192 -0.0085 -0.0182 -0.0149 -0.0149 -0.0121 -0.0318 -0.0085 -0.0149 -0.0211 -0.0144 -0.0085 -0.0085 -0.0152 -0.0085 -0.0155 S

ce i 0.0051 0.0051 0.2055 0.0811 0.1310 0.1310 0.0070 0.0131 0.0312 0.0961 0.1183 0.2451 0.0404 0.0138 Crev -0.0045 -0.0045 -0.0220 -0.0098 -0.0098 -0.0098 -0.0170 -0.0139 -0.0534 -0.0010 -0.0098 -0.0170 -0.0223 -0.0135 -0.0098 -0.0098 -0.0223 -0.0098 Canyon Canyon h g i H 0.0515 0.0515 0.0355 0.0355 0.0752 0.0015 0.0197 0.2907 0.0556 0.0007 0.0649 -0.0079 -0.0079 -0.0129 -0.0253 -0.0065 -0.0274 -0.0220 -0.0220 -0.0129 -0.0066 -0.0003 -0.0127 -0.0127 -0.0149 -0.0204 -0.0127 -0.0127 -0.0339 -0.0127 -0.0244 -0.0033 Plateau Plateau

Upper 0.0330 0.0330 0.0969 0.0969 0.0021 0.1672 0.0168 0.0042 -0.0252 -0.0252 -0.0120 -0.0295 -0.0286 -0.0274 -0.0217 -0.0217 -0.0263 -0.0040 -0.0210 -0.0217 -0.0252 -0.0217 -0.0308 -0.0217 -0.0171 -0.0252 -0.0138 -0.0240 -0.0442 -0.0217 -0.0324 -0.0227 Mesa Top Top Mesa

Lower 0.0018 0.0018 0.1087 0.1614 0.1614 0.1384 0.0002 0.0076 -0.0282 -0.0282 -0.0117 -0.0316 -0.0207 -0.0248 -0.0248 -0.0431 -0.0122 -0.0414 -0.0248 -0.0282 -0.0248 -0.0352 -0.0248 -0.0108 -0.0329 -0.0096 -0.0282 -0.0150 -0.0588 -0.0248 -0.0348 -0.0149 Mesa Top Top Mesa 0.0860 0.0860 0.0293 0.2838 0.0029 0.0029 0.0171 0.0036 0.0063 0.1961 0.0018 -0.0231 -0.0231 -0.0227 -0.0163 -0.0255 -0.0255 -0.0193 -0.0142 -0.0046 -0.0163 -0.0231 -0.0113 -0.0196 -0.0163 -0.0166 -0.0106 -0.0134 -0.0231 -0.0045 -0.0365 -0.0163 d an l Sands Slickrock Slickrock Sands 0.0769 0.0769 0.1187 0.0809 0.0988 0.0988 0.0441 0.1432 0.0552 0.0624 0.0488 0.0210 0.0374 Up -0.0340 -0.0340 -0.0256 -0.0177 -0.0376 -0.0376 -0.0508 -0.0240 -0.0340 -0.0169 -0.0290 -0.0177 -0.0364 -0.0255 -0.0340 -0.0294 -0.0565 -0.0240 -0.0131

y k Roc Slopes Slopes 0.0011 0.0011 0.0124 0.0124 0.0883 0.0754 0.0462 0.0295 0.0119 0.0469 -0.0358 -0.0358 -0.0306 -0.0069 -0.0507 -0.0526 -0.0124 -0.0124 -0.0506 -0.0511 -0.0290 -0.0413 -0.0287 -0.0370 -0.0293 -0.0832 -0.0181 -0.0466 -0.0550 -0.0178 -0.0368 -0.0627 id id Ar 0.0067 0.0067 0.0277 0.0284 0.0039 0.0417 0.0280 0.0337 0.0073 0.1333 0.0152 -0.0089 -0.0089 -0.0224 -0.0001 -0.0355 -0.0115 -0.0610 -0.0114 -0.0114 -0.0947 -0.0162 -0.0119 -0.0198 -0.0469 -0.0260 -0.0110 -0.0260 -0.0331 -0.0488 -0.0110 Lowlands Lowlands d an l Flats Flats 0.0770 0.0770 0.3355 0.3355 0.0993 0.0554 0.0141 0.1470 0.0491 0.0025 -0.0154 -0.0154 -0.0245 -0.0006 -0.0109 -0.0174 -0.0109 -0.0109 -0.0154 -0.0036 -0.0231 -0.0109 -0.0109 -0.0154 -0.0044 -0.0189 -0.0384 -0.0154 -0.0109 -0.0189 -0.0109 -0.0189 Low 2 5 3 1 1 3 2 1 2 1 2 1 7 3 8 2 1 3 1 7 1 6 3 12 40 29 28 Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae 2 Caryophyllaceae Solanaceae Compositae Compositae Berberidaceae Leguminosae Leguminosae Leguminosae Leguminosae Loasaceae Boraginaceae Boraginaceae Compositae Polemoniaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Nyctaginaceae Nyctaginaceae Labiatae Portulacaceae

sp. * 3. Continued. sp. PPENDIX A Muhlenbergia asperifolia Muhlenbergia montana Muhlenbergia pungens Muhlenbergia racemosa Muhlenbergia Muhlenbergia thurberi Muhlenbergia Lychnis drummondii drummondii Lychnis pallidum Lycium grandiflora Lygodesmia canescens Machaeranthera Mahonia repens Medicago lupulina* Melilotus alba* Melilotus officinalis* Melilotus Mentzelia integra arizonica Mertensia fusiformis Mertensia lindleyi Microseris gracilis Microsteris cardinalis Mimulus Mimulus floribundus Mimulus guttatus parryi Mimulus Mirabilis multiflora Mirabilis pumila Monardella odoratissima Montia perfoliata Scientific Name Name Scientific Family # Occ. 68 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.1572 0.1572 0.0022 0.4632 0.1363 0.0020 0.0012 0.0541 0.2472 0.0279 0.0006 0.0407 S -0.0085 -0.0085 -0.0085 -0.0139 -0.0261 -0.1079 -0.0200 -0.0149 -0.0149 -0.0121 -0.0085 -0.0386 -0.0149 -0.0275 -0.0121 -0.0085 -0.0272 -0.0149

ce i 0.0117 0.0117 0.0312 0.1010 0.1121 0.0194 0.0290 0.1087 Crev Canyon Canyon -0.0098 -0.0098 -0.0220 -0.0098 -0.0194 -0.0092 -0.0068 -0.0380 -0.0880 -0.0297 -0.0170 -0.0170 -0.0139 -0.0098 -0.0089 -0.0170 -0.0314 -0.0139 -0.0098 -0.0139 -0.0170 -0.0302 h g i H 0.0122 0.0122 0.0414 0.0197 0.0076 0.0008 0.1617 0.0248 0.0140 0.0071 0.0004 0.0016 Plateau Plateau -0.0127 -0.0127 -0.0058 -0.0001 -0.0284 -0.0179 -0.0497 -0.0893 -0.0291 -0.0220 -0.0220 -0.0085 -0.0220 -0.0235 -0.0179 -0.0058 -0.0220 -0.0371

Upper 0.0645 0.0645 0.0220 0.0278 0.0886 0.1310 0.1029 0.0156 0.0048 0.0015 0.0306 -0.0138 -0.0138 -0.0217 -0.0138 -0.0308 -0.0436 -0.0252 -0.0542 -0.0420 -0.0377 -0.0377 -0.0098 -0.0018 -0.0047 -0.0238 -0.0263 -0.0045 -0.0117 -0.0238 Mesa Top Top Mesa

Lower 0.0014 0.0014 0.0073 0.0235 0.0113 0.0246 0.0515 0.0098 0.2096 0.1849 0.0063 0.0513 0.0150 0.0076 -0.0248 -0.0248 -0.0150 -0.0352 -0.0433 -0.0282 -0.0720 -0.0589 -0.0431 -0.0431 -0.0282 -0.0352 -0.0431 -0.0033 -0.0244 -0.0281 Mesa Top Top Mesa 0.0551 0.0551 0.0019 0.0221 0.0706 0.0685 0.0565 -0.0163 -0.0163 -0.0163 -0.0103 -0.0163 -0.0066 -0.0196 -0.0116 -0.0231 -0.0492 -0.0339 -0.0284 -0.0255 -0.0163 -0.0003 -0.0029 -0.0255 -0.0408 -0.0231 -0.0163 -0.0196 -0.0143 -0.0203 d an l Sands Slickrock Slickrock Sands Up 0.0068 0.0068 0.1056 0.2100 0.0060 0.1714 0.0039 0.0118 0.0220 0.0474 0.0110 0.0411 0.0390 -0.0177 -0.0177 -0.0240 -0.0051 -0.0240 -0.0290 -0.0340 -0.0750 -0.0381 -0.0339 -0.0089 -0.0358 -0.0339 -0.0249 -0.0157 -0.0174 -0.0067

y k Roc Slopes Slopes 0.0019 0.0019 0.0777 0.0494 0.0095 0.1969 0.0187 0.0088 0.0039 0.0236 0.0315 0.0142 0.0154 0.0177 0.0152 0.0249 0.0457 0.1760 -0.0368 -0.0368 -0.0351 -0.0138 -0.0370 -0.0413 -0.0325 -0.0459 -0.0429 -0.0118 -0.0199 -0.0321 id id Ar 0.0081 0.0081 0.2280 0.0051 0.0449 0.0721 0.0719 0.0666 0.0005 0.0004 0.0296 -0.0068 -0.0068 -0.0204 -0.0303 -0.0001 -0.0030 -0.0198 -0.0057 -0.0220 -0.0026 -0.0254 -0.0258 -0.1407 -0.0052 -0.0305 -0.0008 -0.0539 -0.0216 -0.0032 Lowlands Lowlands d an l Flats Flats 0.0225 0.0225 0.0512 0.1635 0.1030 0.0158 0.0226 0.0069 0.0017 -0.0109 -0.0109 -0.0109 -0.0033 -0.0154 -0.0224 -0.0068 -0.0871 -0.0189 -0.0154 -0.0109 -0.0154 -0.0060 -0.0573 -0.0154 -0.0194 -0.0109 -0.0262 -0.0269 -0.0189 -0.0439 Low 1 1 5 1 9 2 2 9 3 3 2 1 2 4 2 3 2 1 6 2 3 12 17 54 10 38 23 120 120 Ranunculaceae Cruciferae Polemoniaceae Solanaceae Onagraceae Onagraceae Onagraceae Onagraceae Cactaceae Cactaceae Cactaceae Cactaceae Cactaceae Orobanchaceae Orobanchaceae Gramineae Umbelliferae Celastraceae Gramineae Boraginaceae Scrophulariaceae Leguminosae Polypodiaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae

3. Continued.

sp. sp. PPENDIX A Scientific Name Name Scientific Family # Occ. Myosurus cupulatus officinale Nasturtium breweri Navarretia Nicotiana attenuata caespitosa Oenothera longissima Oenothera pallida Oenothera Oenothera erinacea Opuntia macrorhiza Opuntia Opuntia phaeacantha Opuntia polyacantha whipplei Opuntia fasciculata Orobanche uniflora Orobanche micrantha Oryzopsis occidentalis Osmorhiza myrsinites Pachystima Panicum setosa Pectocarya centranthera Pedicularis mephiticum Pediomelum Pellaea truncata Penstemon angustifolius Penstemon barbatus comarrhenus Penstemon Penstemon confusus Penstemon eatonii 2015] PLANT COALITION GROUPS OF ZION NP 69

bank bank tream- 0.0380 0.0380 0.0050 0.0088 0.1188 0.0094 0.0047 0.3451 0.0376 0.0114 S -0.0124 -0.0124 -0.0438 -0.0456 -0.0167 -0.0085 -0.0184 -0.0275 -0.0192 -0.0121 -0.0211 -0.0085 -0.0675 -0.0234 -0.0327 -0.0149 -0.0149 -0.0229 -0.0149 -0.0085

ce i Crev 0.0411 0.0411 0.1111 0.0277 0.1274 0.0676 0.2350 0.0143 0.0002 Canyon Canyon -0.0099 -0.0099 -0.0342 -0.0360 -0.0098 -0.0295 -0.0293 -0.0220 -0.0139 -0.0242 -0.0098 -0.0098 -0.0503 -0.0293 -0.0335 -0.0044 -0.0170 -0.0170 -0.0261 -0.0170 -0.0098 h g i H 0.0924 0.0924 0.0331 0.0460 0.0165 0.0069 0.1026 0.0565 Plateau Plateau -0.0374 -0.0374 -0.0542 -0.0127 -0.0468 -0.0127 -0.0254 -0.0130 -0.0313 -0.0032 -0.0127 -0.0512 -0.0028 -0.0264 -0.0079 -0.0226 -0.0127 -0.0220 -0.0137 -0.0279 -0.0143 -0.0065

Upper 0.0720 0.0720 0.2105 0.2389 0.0762 0.0433 0.2187 0.0399 0.0155 0.0418 0.2013 -0.0125 -0.0125 -0.0335 -0.0217 -0.0480 -0.0091 -0.0217 -0.0489 -0.0243 -0.0365 -0.0256 -0.0138 -0.0217 -0.0116 -0.0308 -0.0346 -0.0138 -0.0113 -0.0125 Mesa Top Top Mesa

Lower 0.0574 0.0574 0.0904 0.0101 0.1847 0.0020 0.0352 0.0491 0.1193 0.0484 0.0003 0.0871 0.1630 0.0474 -0.0007 -0.0007 -0.0416 -0.0248 -0.0624 -0.0248 -0.0559 -0.0060 -0.0399 -0.0231 -0.0150 -0.0248 -0.0352 -0.0150 -0.0187 -0.0187 Mesa Top Top Mesa 0.2021 0.2021 0.0846 0.1297 0.0041 0.0074 0.3143 0.0148 0.1294 0.0721 0.0010 -0.0180 -0.0180 -0.0471 -0.0163 -0.0310 -0.0351 -0.0311 -0.0354 -0.0056 -0.0163 -0.0282 -0.0285 -0.0196 -0.0332 -0.0163 -0.0166 -0.0154 -0.0416 -0.0168 d an l Sands Slickrock Slickrock Sands Up 0.1052 0.1052 0.2316 0.0259 0.0268 0.0973 0.1886 0.0545 0.1120 0.1040 -0.0317 -0.0317 -0.0284 -0.0034 -0.0440 -0.0137 -0.0263 -0.0240 -0.0511 -0.0356 -0.0094 -0.0177 -0.0246 -0.0061 -0.0290 -0.0285 -0.0240 -0.0097 -0.0185 -0.0163

y k Roc Slopes Slopes 0.0790 0.0790 0.0773 0.0043 0.0837 0.0496 0.0457 0.0028 0.0044 0.1251 0.0293 0.1106 0.0859 0.0691 0.0068 0.0326 -0.0169 -0.0169 -0.0164 -0.0258 -0.0368 -0.0140 -0.0066 -0.0582 -0.0293 -0.0154 -0.0409 -0.0355 -0.0215 -0.0141 id id Ar 0.0759 0.0759 0.1018 0.1095 0.1215 0.0051 0.0417 0.0037 0.0209 -0.0318 -0.0318 -0.0565 -0.0162 -0.0437 -0.0415 -0.0473 -0.0204 -0.0290 -0.0612 -0.0651 -0.0600 -0.0520 -0.0238 -0.0010 -0.0087 -0.0088 -0.0200 -0.0018 -0.0058 -0.0161 Lowlands Lowlands d an l Flats Flats 0.0007 0.0007 0.0215 0.0128 0.0196 0.0554 0.0053 0.0857 0.1195 -0.0245 -0.0245 -0.0593 -0.0532 -0.0109 -0.0032 -0.0179 -0.0016 -0.0109 -0.0385 -0.0144 -0.0498 -0.0248 -0.0767 -0.0331 -0.0154 -0.0049 -0.0189 -0.0106 -0.0189 -0.0109 Low 5 1 1 5 2 6 1 1 2 8 1 3 3 7 3 1 27 26 19 30 10 12 42 10 54 11 14 Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Rosaceae Compositae Leguminosae Compositae Rosaceae Hydrophyllaceae Hydrophyllaceae Hydrophyllaceae Hydrophyllaceae Saxifragaceae Polemoniaceae Polemoniaceae Viscaceae Gramineae Solanaceae Solanaceae Cruciferae Cruciferae Cruciferae Cruciferae Cruciferae Scrophulariaceae 13 Scrophulariaceae 3. Continued. PPENDIX A Scientific Name Name Scientific Family # Occ. Penstemon higginsii higginsii Penstemon humilis Penstemon laevis Penstemon linarioides Penstemon pachyphyllus Penstemon palmeri Penstemon rostriflorus Penstemon ramosissimum Peraphyllum tenella Perityle Peteria thompsoniae Petradoria pumila caespitosum Petrophytum fremontii Phacelia heterophylla Phacelia ivesiana Phacelia palmeri Phacelia microphyllus Philadelphus austromontana Phlox Phlox longifolia juniperinum Phoradendron australis Phragmites hederifolia Physalis Physalis longifolia chambersii Physaria ludoviciana Physaria newberryi Physaria rectipes Physaria wardii Physaria 70 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.0094 0.0094 0.2434 0.2027 0.0228 0.3936 0.0664 0.0201 0.0212 0.0044 0.1094 0.3576 -0.0085 -0.0085 -0.0640 -0.0597 -0.0322 -0.0315 -0.0181 -0.0837 -0.0172 -0.0085 -0.0010 -0.0289 -0.0121 -0.0339 -0.0475 -0.0480 -0.0524 -0.0085 S

ce i 0.0285 0.0285 0.0051 0.0391 0.2726 0.0149 0.0426 0.0006 0.2559 0.1626 0.0224 0.0666 0.0371 Crev Canyon Canyon -0.0030 -0.0030 -0.0481 -0.0790 -0.0360 -0.0280 -0.0016 -0.0072 -0.0162 -0.0098 -0.0403 -0.0146 -0.0330 -0.0388 -0.0563 -0.0597 -0.0098 h g i H 0.0583 0.0583 0.2466 0.1711 0.0408 0.0008 0.0909 0.2923 0.0309 0.0407 0.0241 0.1418 0.0263 Plateau Plateau -0.0127 -0.0127 -0.0540 -0.0808 -0.0050 -0.0292 -0.0329 -0.0127 -0.0417 -0.0207 -0.0127 -0.0255 -0.0428 -0.0468 -0.0297 -0.0805 -0.0127

Upper 0.0135 0.0135 0.1513 0.0717 0.0891 0.0855 0.0676 0.0042 0.1832 0.1804 -0.0930 -0.0930 -0.0376 -0.0593 -0.0217 -0.0221 -0.0258 -0.0227 -0.0217 -0.0263 -0.0487 -0.0116 -0.0337 -0.0660 -0.0232 -0.0126 -0.0317 -0.0825 -0.0138 -0.0217 Mesa Top Top Mesa

Lower 0.0473 0.0473 0.2160 0.0611 0.0577 0.0050 0.0307 0.0553 0.1318 0.1792 -0.1054 -0.1054 -0.0564 -0.0675 -0.0248 -0.0307 -0.0455 -0.0237 -0.0248 -0.0031 -0.0613 -0.0199 -0.0449 -0.0747 -0.0009 -0.0007 -0.0049 -0.0980 -0.0150 -0.0248 Mesa Top Top Mesa 0.0168 0.0168 0.1993 0.1186 0.0205 0.0311 0.0592 0.1328 0.0421 0.0422 0.0036 -0.0519 -0.0519 -0.1071 -0.0229 -0.0388 -0.0113 -0.0183 -0.0409 -0.0163 -0.0362 -0.0095 -0.0332 -0.0303 -0.0532 -0.0071 -0.0602 -0.0355 -0.0280 -0.0113 d an l Sands Slickrock Slickrock Sands 0.0150 0.0150 0.1735 0.0399 0.1283 0.0318 0.1548 0.0472 0.0959 0.0354 0.0456 0.0488 Up -0.0240 -0.0240 -0.0616 -0.0542 -0.0169 -0.0320 -0.0276 -0.0177 -0.0402 -0.0534 -0.0100 -0.0404 -0.0183 -0.0232 -0.0679 -0.0027 -0.0691 -0.0169

y k Roc Slopes Slopes 0.0064 0.0064 0.0969 0.1964 0.2132 0.0621 0.0239 0.0437 0.0241 0.2131 0.0295 -0.0675 -0.0675 -0.0524 -0.0240 -0.0368 -0.0550 -0.0991 -0.0321 -0.0363 -0.0756 -0.0359 -0.0766 -0.0415 -0.0447 -0.0107 -0.0371 -0.0137 -0.0259 -0.0232 id id Ar 0.1617 0.1617 0.1362 0.1065 0.0257 0.0173 0.2031 0.0794 0.1136 0.0337 -0.0204 -0.0204 -0.0799 -0.1236 -0.0162 -0.0296 -0.0620 -0.0598 -0.0150 -0.0693 -0.0470 -0.0177 -0.0290 -0.0582 -0.0849 -0.0327 -0.0204 -0.0642 -0.2123 -0.0117 Lowlands Lowlands d an l Flats Flats 0.1238 0.1238 0.0487 0.0848 0.0185 0.0196 0.0293 0.0437 0.0240 -0.0109 -0.0109 -0.0363 -0.0639 -0.0574 -0.1391 -0.0045 -0.0219 -0.0015 -0.0269 -0.0139 -0.0154 -0.0476 -0.0189 -0.0154 -0.0140 -0.0398 -0.0835 -0.0617 -0.0109 -0.0109 Low 1 8 1 3 4 1 6 6 2 4 3 2 1 1 53 64 60 13 14 10 18 11 15 36 50 190 190 135 Rosaceae Pinaceae Pinaceae Pinaceae Plantaginaceae Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae Polypodiaceae Salicaceae Salicaceae Rosaceae Pinaceae Leguminosae Leguminosae Polypodiaceae Monotropaceae Rosaceae Rosaceae Fagaceae Fagaceae Ranunculaceae Ranunculaceae Polygonaceae 39

3. Continued. sp.

sp. PPENDIX A Scientific Name Name Scientific Family # Occ. Physocarpus alternans alternans Physocarpus Pinus edulis Pinus monophylla Pinus ponderosa Plantago patagonica Poa bigelovii Poa bulbosa* compressa* Poa fendleriana Poa pratensis* Poa secunda Poa Poa Polygonum douglasii hesperium Polypodium fremontii Populus tremuloides Populus Prunus virginiana Pseudotsuga menziesii Psoralidium lanceolatum fremontii Psorothamnus Pteridium aquilinum andromedea Pterospora mexicana Purshia tridentata Purshia gambelii Quercus Quercus turbinella Ranunculus andersonii Ranunculus 2015] PLANT COALITION GROUPS OF ZION NP 71

bank bank tream- 0.0057 0.0057 0.0255 0.0438 0.0114 0.0432 0.4594 0.0246 0.1572 0.1572 0.1908 0.0600 0.0600 0.1572 0.0200 0.0034 -0.0388 -0.0388 -0.0085 -0.0121 -0.0149 -0.0085 -0.0121 -0.0025 -0.0149 -0.0180 -0.0121 -0.0085 -0.0606 -0.0606 -0.0271 -0.0192 S

ce i 0.0501 0.0501 0.0002 0.2611 0.0332 0.0423 0.0117 0.0390 0.0117 0.0131 0.0922 0.0031 0.1728 0.0264 0.0063 0.0063 0.0117 0.1054 Crev -0.0139 -0.0139 -0.0425 -0.0035 -0.0098 -0.0139 -0.0170 -0.0139 -0.0043 Canyon Canyon -0.0054 -0.0054 -0.0397 -0.0131 -0.0171 h g i H 0.0826 0.0826 0.2403 0.0373 0.0088 0.0239 0.0122 0.0122 0.0304 0.0099 0.3109 0.3109 0.0122 0.0085 -0.0179 -0.0179 -0.0439 -0.0058 -0.0127 -0.0179 -0.0220 -0.0029 -0.0127 -0.0179 -0.0095 -0.0129 -0.0032 -0.0288 -0.0288 -0.0453 -0.0178 -0.0124 Plateau Plateau

Upper 0.0130 0.0130 0.0501 0.0042 0.0038 0.0038 0.0478 -0.0308 -0.0308 -0.0392 -0.0059 -0.0138 -0.0310 -0.0308 -0.0377 -0.0182 -0.0273 -0.0217 -0.0092 -0.0217 -0.0217 -0.0252 -0.0005 -0.0178 -0.0260 -0.0242 -0.0217 -0.0217 -0.0629 -0.0547 -0.0278 -0.0040 Mesa Top Top Mesa

Lower 0.0722 0.0722 0.0053 0.0094 0.0076 0.0662 0.0662 0.0048 -0.0352 -0.0352 -0.0629 -0.0051 -0.0051 -0.0150 -0.0278 -0.0352 -0.0431 -0.0166 -0.0301 -0.0248 -0.0248 -0.0248 -0.0282 -0.0060 -0.0182 -0.0269 -0.0125 -0.0125 -0.0248 -0.0538 -0.0523 -0.0307 Mesa Top Top Mesa 0.0088 0.0088 0.0898 0.1569 0.1751 0.2741 0.1932 0.0036 0.1136 0.1136 0.0257 0.0129 0.0094 -0.0231 -0.0231 -0.0489 -0.0079 -0.0163 -0.0177 -0.0163 -0.0231 -0.0226 -0.0185 -0.0249 -0.0163 -0.0163 -0.0113 -0.0231 -0.0284 -0.0284 -0.0163 -0.0565 d an l Sands Slickrock Slickrock Sands 0.0311 0.0311 0.1417 0.0248 0.0623 0.0405 0.1919 0.1919 0.0456 0.0642 0.0450 Up -0.0340 -0.0340 -0.0483 -0.0175 -0.0149 -0.0490 -0.0240 -0.0420 -0.0340 -0.0335 -0.0340 -0.0148 -0.0240 -0.0240 -0.0169 -0.0340 -0.0009 -0.0312 -0.0312 -0.0240 -0.0524

y k Roc Slopes Slopes 0.1113 0.1113 0.0011 0.0022 0.0290 0.0336 0.1615 0.1615 0.1785 0.0061 0.0558 -0.0522 -0.0522 -0.0878 -0.0215 -0.0585 -0.0411 -0.0469 -0.0466 -0.0368 -0.0006 -0.0368 -0.0287 -0.0464 -0.0064 -0.0156 -0.0063 -0.0142 -0.0552 -0.0552 -0.0368 -0.0308 id id Ar 0.1709 0.1709 0.0029 0.0037 0.0270 0.0757 0.0039 0.1196 0.1196 0.0432 -0.0166 -0.0166 -0.0161 -0.0648 -0.0390 -0.0139 -0.0290 -0.0204 -0.0159 -0.0204 -0.0197 -0.0290 -0.0245 -0.0703 -0.0255 -0.0015 -0.0314 -0.0314 -0.0204 -0.1087 -0.0684 -0.0317 Lowlands Lowlands d an l Flats Flats 0.0241 0.0241 0.0277 0.1195 0.0912 -0.0011 -0.0011 -0.0109 -0.0109 -0.0219 -0.0189 -0.0154 -0.0189 -0.0109 -0.0109 -0.0109 -0.0109 -0.0154 -0.0154 -0.0189 -0.0462 -0.0154 -0.0109 -0.0189 -0.0189 -0.0109 -0.0802 -0.0242 -0.0160 -0.0189 -0.0245 Low 2 1 1 1 4 2 3 2 3 1 1 1 1 2 2 3 2 1 1 3 5 23 12 17 99 26 17 Ranunculaceae Anacardiaceae Saxifragaceae Leguminosae Rosaceae Rubiaceae Rosaceae Polygonaceae Salicaceae Salicaceae Labiatae Primulaceae Saxifragaceae Cyperaceae Cyperaceae Gramineae Crassulaceae Crassulaceae Selaginellaceae Selaginellaceae Selaginellaceae Compositae Compositae Compositae Compositae Elaeagnaceae Caryophyllaceae Caryophyllaceae Compositae 3 Compositae 3. Continued. PPENDIX A Ranunculus testiculatus* Rhus aromatica velutinum Ribes neomexicana Robinia Rosa woodsii Rubia tinctoria* leucodermis Rubus hymenosepalus Rumex Salix exigua scouleriana Salix Salvia dorrii Samolus floribundus rhomboidea Saxifraga microcarpus Scirpus pungens Scirpus Secale cereale* Sedum debile Sedum lanceolatum underwoodii Selaginella Selaginella utahensis Selaginella watsonii Scientific Name Name Scientific Family # Occ. Senecio eremophilus eremophilus Senecio Senecio hydrophilus Senecio multilobatus Senecio spartioides Shepherdia rotundifolia antirrhina Silene verecunda Silene 72 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.0090 0.0090 0.2472 0.0089 0.3652 0.0469 0.0665 0.0380 0.0318 0.0320 -0.0060 -0.0060 -0.0085 -0.0085 -0.0134 -0.0121 -0.0224 -0.0085 -0.0061 -0.0121 -0.0085 -0.0121 -0.0371 -0.0490 -0.0355 -0.0085 -0.0121 -0.0292 -0.0234 -0.0533 S

ce i 0.0194 0.0194 0.2531 0.0379 0.1476 0.0915 0.1957 0.0534 0.0698 Crev -0.0197 -0.0197 -0.0098 -0.0098 -0.0139 -0.0347 -0.0098 -0.0068 -0.0588 -0.0139 -0.0098 -0.0100 -0.0602 -0.0721 -0.0184 -0.0069 -0.0098 -0.0139 -0.0332 -0.0243 -0.0623 Canyon Canyon h g i H 0.0272 0.0272 0.0097 0.0463 0.0284 0.0889 0.0413 0.0187 0.3314 0.2524 0.2604 0.0254 -0.0127 -0.0127 -0.0127 -0.0036 -0.0423 -0.0127 -0.0179 -0.0708 -0.0179 -0.0127 -0.0179 -0.0463 -0.0122 -0.0788 -0.0127 -0.0467 -0.0386 -0.0617 Plateau Plateau

Upper 0.1036 0.1036 0.0530 0.0497 0.0911 0.0612 0.0987 0.0374 -0.0222 -0.0222 -0.0308 -0.0148 -0.0142 -0.0217 -0.0217 -0.0332 -0.0643 -0.0217 -0.0332 -0.0252 -0.0813 -0.0308 -0.0217 -0.0196 -0.0537 -0.0169 -0.0408 -0.0217 -0.0779 -0.0412 Mesa Top Top Mesa

Lower 0.0248 0.0248 0.0102 0.0614 0.0068 0.0271 0.0173 0.0865 0.0734 -0.0350 -0.0350 -0.0352 -0.0248 -0.0248 -0.0207 -0.0771 -0.0248 -0.0374 -0.0282 -0.1019 -0.0352 -0.0248 -0.0212 -0.0459 -0.0597 -0.0027 -0.0135 -0.0248 -0.0881 -0.0483 Mesa Top Top Mesa 0.0917 0.0917 0.1890 0.0013 0.0001 0.2097 0.0009 -0.0249 -0.0249 -0.0196 -0.0215 -0.0163 -0.0163 -0.0231 -0.0565 -0.0163 -0.0231 -0.0302 -0.0142 -0.0308 -0.0163 -0.0153 -0.0626 -0.0549 -0.0363 -0.0285 -0.0100 -0.0155 -0.0373 -0.0797 d an l Sands Slickrock Slickrock Sands 0.0046 0.0046 0.1236 0.1172 0.0637 0.0327 0.0077 0.0203 Up -0.0328 -0.0328 -0.0290 -0.0427 -0.0334 -0.0123 -0.0240 -0.0338 -0.0142 -0.0691 -0.0115 -0.0417 -0.0340 -0.0340 -0.0712 -0.0115 -0.0144 -0.0170 -0.0215 -0.0240 -0.0627 -0.0189

y k Roc Slopes Slopes 0.0237 0.0237 0.0846 0.0299 0.0886 0.0749 0.1070 0.1502 -0.0258 -0.0258 -0.0425 -0.0248 -0.0640 -0.0368 -0.0368 -0.0889 -0.0428 -0.0176 -0.0075 -0.0264 -0.0550 -0.0522 -0.0303 -0.1064 -0.0264 -0.0105 -0.0723 -0.0550 -0.0135 -0.0200 id id Ar 0.0076 0.0076 0.0024 0.0710 0.0424 0.0677 0.0213 0.1120 0.0813 0.0040 0.1855 0.1109 0.0388 -0.0488 -0.0488 -0.0355 -0.0162 -0.0162 -0.1190 -0.0280 -0.0100 -0.0043 -0.0331 -0.0230 -0.0822 -0.0043 -0.0310 -0.0631 -0.0389 -0.0227 Lowlands Lowlands d an l Flats Flats 0.0640 0.0640 0.0271 0.0085 0.2053 0.0188 0.0570 0.0823 0.0048 0.0315 0.0881 0.0436 0.0249 0.0411 -0.0139 -0.0139 -0.0189 -0.0109 -0.0109 -0.0006 -0.0154 -0.0109 -0.0189 -0.0109 -0.0564 -0.0221 -0.0077 -0.0254 -0.0200 -0.0361 Low 4 2 6 3 1 1 3 2 1 3 2 2 1 2 5 1 2 38 14 50 18 51 39 65 12 15 11 38 Cruciferae Cruciferae Cruciferae Liliaceae Liliaceae Solanaceae Solanaceae Compositae Compositae Malvaceae Malvaceae Malvaceae Compositae Gramineae Gramineae Cruciferae Caryophyllaceae Compositae Compositae Compositae Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae Gramineae Cruciferae Cruciferae

3. Continued. * sp. sp. PPENDIX A Sisymbrium altissimum* altissimum* Sisymbrium irio* Sisymbrium racemosa Smilacina Smilacina stellata sarrachoides* Solanum Solanum velutina Solidago Sonchus ambigua Sphaeralcea grossulariifolia Sphaeralcea parvifolia Sphaeralcea ruthiae Sphaeromeria contractusSporobolus cryptandrusSporobolus Stanleya pinnata jamesiana Stellaria exigua Stephanomeria Stephanomeria runcinata tenuifolia Stephanomeria comata Stipa Stipa hymenoides lettermanii Stipa Stipa nelsonii neomexicana Stipa Stipa pinetorum Stipa speciosa Streptanthella longirostris Streptanthus cordatus Scientific Name Name Scientific Family # Occ. 2015] PLANT COALITION GROUPS OF ZION NP 73

bank bank tream- 0.4269 0.4269 0.1004 0.0461 0.0474 0.0308 0.0144 0.0538 0.0077 0.0310 0.0045 0.2472 0.1572 0.3743 0.0130 -0.0085 -0.0085 -0.0289 S -0.0289 -0.0289 -0.0121 -0.0198 -0.0085 -0.0260 -0.0116 -0.0085 -0.0383 -0.0085 -0.0085 -0.0212

ce i 0.0266 0.0266 0.2364 0.2305 0.1166 0.0169 0.1317 0.0265 0.0194 0.0117 0.0353 0.1323 0.0347 Crev Canyon Canyon -0.0098 -0.0098 -0.0330 -0.0213 -0.0213 -0.0139 -0.0018 -0.0098 -0.0185 -0.0253 -0.0098 -0.0098 -0.0429 -0.0101 -0.0068 -0.0098 -0.0243 h g i H 0.0332 0.0332 0.2005 0.0066 0.2013 0.1990 0.0083 0.0581 0.0007 0.0161 0.0961 0.0456 0.0097 0.0122 0.1451 0.2497 0.0675 Plateau Plateau -0.0127 -0.0127 -0.0428 -0.0179 -0.0179 -0.0055 -0.0027 -0.0296 -0.0127 -0.0375 -0.0179 -0.0127 -0.0127

Upper 0.1982 0.1982 0.1700 0.0013 0.0435 0.0085 0.0434 0.0034 0.1074 0.0916 -0.0037 -0.0037 -0.0527 -0.0196 -0.0196 -0.0045 -0.0489 -0.0108 -0.0048 -0.0249 -0.0401 -0.0098 -0.0119 -0.0030 -0.0540 -0.0308 -0.0167 -0.0217 -0.0374 -0.0217 Mesa Top Top Mesa

Lower 0.1911 0.1911 0.1381 0.0072 0.0401 0.0191 0.0806 0.0174 -0.0023 -0.0023 -0.0511 -0.0212 -0.0212 -0.0033 -0.0559 -0.0170 -0.0331 -0.0327 -0.0466 -0.0248 -0.0248 -0.0053 -0.0627 -0.0355 -0.0352 -0.0044 -0.0282 -0.0248 -0.0340 -0.0248 Mesa Top Top Mesa 0.0441 0.0441 0.0331 0.0290 0.0139 0.0103 0.2031 0.1027 -0.0104 -0.0104 -0.0533 -0.0251 -0.0251 -0.0140 -0.0478 -0.0322 -0.0134 -0.0028 -0.0163 -0.0083 -0.0108 -0.0173 -0.0323 -0.0196 -0.0196 -0.0163 -0.0105 -0.0499 -0.0163 -0.0318 d an l Sands Slickrock Slickrock Sands Up 0.1325 0.1325 0.0890 0.0645 0.1042 0.2178 0.0103 -0.0112 -0.0112 -0.0462 -0.0212 -0.0212 -0.0045 -0.0492 -0.0476 -0.0492 -0.0473 -0.0046 -0.0139 -0.0240 -0.0340 -0.0421 -0.0290 -0.0228 -0.0240 -0.0092 -0.0372 -0.0652 -0.0177 -0.0051

y k Roc Slopes Slopes 0.0285 0.0285 0.0796 0.0466 0.0466 0.0363 0.0630 0.0414 0.0578 -0.0337 -0.0337 -0.0200 -0.0154 -0.0142 -0.0696 -0.0846 -0.0848 -0.0546 -0.0290 -0.0414 -0.0837 -0.0522 -0.0552 -0.0072 -0.0368 -0.0217 -0.0698 -0.0621 -0.0107 -0.0190 id id Ar 0.0045 0.0045 0.1751 0.0028 0.0028 0.0346 0.0220 0.0149 0.0550 -0.0538 -0.0538 -0.1199 -0.0111 -0.0253 -0.0636 -0.0212 -0.0697 -0.0363 -0.0411 -0.0106 -0.0260 -0.0352 -0.0230 -0.0608 -0.0161 -0.0204 -0.0204 -0.0370 -0.0951 -0.0363 Lowlands Lowlands d an l Flats Flats 0.0097 0.0097 0.2444 0.1011 0.0196 0.0440 -0.0109 -0.0109 -0.0219 -0.0245 -0.0245 -0.0154 -0.0154 -0.0109 -0.0245 -0.0062 -0.0057 -0.0291 -0.0219 -0.0252 -0.0109 -0.0109 -0.0154 -0.0477 -0.0014 -0.0154 -0.0109 -0.0109 -0.0103 -0.0117 Low 1 2 1 5 9 7 4 1 1 2 8 2 2 1 1 5 1 11 58 11 12 10 22 10 43 12 Liliaceae Liliaceae Gentianaceae Gentianaceae Gentianaceae Caprifoliaceae Portulacaceae Tamaricaceae Compositae Compositae Ranunculaceae Cruciferae Leguminosae Cruciferae Compositae Compositae Anacardiaceae Commelinaceae Compositae Zygophyllaceae Leguminosae Leguminosae Typhaceae Valerianaceae Scrophulariaceae Leguminosae Leguminosae Compositae Gentianaceae 11 Gentianaceae

3. Continued. sp. PPENDIX A Scientific Name Name Scientific Family # Occ. Streptopus amplexifolius amplexifolius Streptopus albomarginata Swertia Swertia radiata radiata Swertia utahensis Swertia oreophilus Symphoricarpos Talinum parviflorum chinensis* Tamarix officinale* Taraxacum canescens Tetradymia Thalictrum fendleri Thelypodium laxiflorum montana Thermopsis curvipes Thysanocarpus incana Townsendia montana Townsendia rydbergii Toxicodendron occidentalis Tradescantia Tragopogon dubius* terrestris* Tribulus Trifolium longipes Trifolium Typha latifolia Valeriana occidentalis thapsus* Verbascum americana Vicia ludoviciana Vicia multiflora Viguiera 74 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

bank bank tream- 0.0892 0.0892 0.0255 0.0709 0.3576 -0.0275 -0.0275 -0.0085 -0.0192 -0.0521 -0.0318 -0.0039 -0.0302 S

ce i 0.3300 0.3300 0.0023 0.1491 0.0878 0.0371 0.1956 -0.0314 -0.0314 -0.0192 -0.0344 -0.0392 -0.0320 Crev Canyon Canyon h g i H 0.0734 0.0734 0.0934 0.0136 0.0263 0.0217 -0.0322 -0.0322 -0.0004 -0.0253 -0.0629 -0.0489 -0.0016 Plateau Plateau

Upper 0.1282 0.1282 0.1945 -0.0347 -0.0347 -0.0040 -0.0138 -0.0138 -0.0292 -0.0217 -0.0226 -0.0588 -0.0293 Mesa Top Top Mesa

Lower 0.2702 0.2702 0.0140 0.0843 -0.0120 -0.0120 -0.0150 -0.0150 -0.0313 -0.0248 -0.0337 -0.0682 -0.0038 Mesa Top Top Mesa 0.0225 0.0225 0.0567 0.1696 0.0550 0.2515 -0.0163 -0.0163 -0.0446 -0.0163 -0.0113 -0.0344 -0.0432 d an l Sands Slickrock Slickrock Sands 0.0023 0.0023 0.0301 0.1379 0.0005 Up -0.0388 -0.0388 -0.0149 -0.0334 -0.0240 -0.0169 -0.0227 -0.0498

y k Roc Slopes Slopes 0.0052 0.0052 0.1162 0.0981 -0.0655 -0.0655 -0.0368 -0.0220 -0.0290 -0.0428 -0.0232 -0.0283 -0.0120 id id Ar 0.1913 0.1913 -0.0441 -0.0441 -0.0133 -0.0474 -0.0162 -0.0162 -0.0350 -0.0117 -0.0505 -0.0558 -0.0079 Lowlands Lowlands d an l Flats Flats 0.0039 0.0039 0.1141 -0.0245 -0.0245 -0.0109 -0.0109 -0.0109 -0.0245 -0.0109 -0.0660 -0.0066 -0.0160 Low 5 1 1 1 5 1 10 41 17 15 12 Violaceae Violaceae Violaceae Violaceae Vitaceae Polypodiaceae Compositae Compositae Agavaceae Agavaceae Onagraceae Liliaceae 3. Continued. PPENDIX A Viola canadensis Viola nuttallii purpurea Viola arizonica Vitis Woodsia oregana Wyethia arizonica strumarium Xanthium Yucca angustissima baccata Yucca Zauschneria latifolia Zigadenus paniculatus Scientific Name Name Scientific Family # Occ. *Exotic species (Welsh et al. 2003, Fertig and Alexander 2009) and Alexander et al. 2003, Fertig species (Welsh *Exotic 2015] PLANT COALITION GROUPS OF ZION NP 75 0.0262 0.0262 0.0077 0.0185 0.0031 0.1363 bank bank -0.0065 -0.0065 -0.0061 -0.0211 -0.0260 -0.0138 -0.0275 -0.0149 -0.0224 -0.0315 -0.0584 -0.0544 -0.0086 -0.0469 -0.0510 -0.0639 -0.0189 -0.0339 -0.0160 -0.0214 -0.0374 -0.0355 Stream- Stream- ce ce i 0.0243 0.0243 -0.0873 -0.0873 -0.0427 -0.0588 -0.0101 -0.0282 -0.0297 -0.0393 -0.0260 -0.0314 -0.0170 -0.0347 -0.0360 -0.0969 -0.0642 -0.0962 -0.0819 -0.0675 -0.0776 -0.0615 -0.0369 -0.0261 -0.0520 -0.0394 -0.0721 -0.0092 Crev Canyon Canyon

h g i 0.0961 0.0961 0.0166 0.0116 H -0.1083 -0.1083 -0.0518 -0.0708 -0.0407 -0.0385 -0.0646 -0.0225 -0.0407 -0.0220 -0.0423 -0.0466 -0.1140 -0.0865 -0.1043 -0.0946 -0.0683 -0.0746 -0.0737 -0.0503 -0.0302 -0.0554 -0.0788 -0.0284 Plateau Plateau s to coalition groups (columns; see also 0.0045 0.0045 0.0085 -0.1512 -0.1512 -0.0813 -0.0609 -0.0387 -0.0594 -0.0888 -0.0178 -0.0634 -0.0263 -0.0643 -0.0800 -0.0963 -0.1239 -0.1114 -0.1123 -0.1138 -0.0796 -0.0745 -0.0779 -0.0550 -0.0588 -0.0854 -0.0408 -0.0436 Upper Upper Mesa Top Top Mesa tis 1959). Affinities are mean pairwise association tis 1959). Affinities rder of percent occurrence within vegetation types. 0.0486 0.0486 -0.1832 -0.1832 -0.1019 -0.0355 -0.0766 -0.0521 -0.0755 -0.0986 -0.0208 -0.0797 -0.0431 -0.0771 -0.0914 -0.1116 -0.1439 -0.1590 -0.1467 -0.1262 -0.1192 -0.0816 -0.0797 -0.0626 -0.0556 -0.0888 -0.0027 -0.0433 Lower Lower Mesa Top Top Mesa -0.1214 -0.1214 -0.0683 -0.0302 -0.0323 -0.0488 -0.0233 -0.0475 -0.0026 -0.0291 -0.0447 -0.0255 -0.0565 -0.0601 -0.1029 -0.0855 -0.0984 -0.0069 -0.0871 -0.0596 -0.0704 -0.0476 -0.0395 -0.0819 -0.0630 -0.0549 -0.0116

d an l 0.1236 0.1236 0.0515 0.0993 0.0211 0.0327 0.1056 -0.0590 -0.0590 -0.0601 -0.0421 -0.0748 -0.0662 -0.0709 -0.0051 -0.0069 -0.0708 -0.0376 -0.0691 -0.0830 -0.0990 -0.0696 -0.1195 -0.0141 -0.0719 -0.0562 -0.0494 -0.0872 Sands Slickrock Slickrock Sands Up y k 0.0710 0.0710 0.0237 0.0011 0.1513 0.0731 0.0670 0.0657 0.0402 0.1317 0.0867 0.0316 0.0713 0.0091 0.0886 0.0494 -0.0514 -0.0514 -0.0837 -0.0910 -0.0836 -0.0609 -0.0022 -0.0649 -0.0506 -0.0075 -0.0393 -0.0365 Roc Slopes Slopes

id 0.2135 0.2135 0.0081 0.0424 0.1277 0.2167 0.1291 0.0716 0.0067 0.0710 0.2448 0.2558 0.3680 0.2817 0.2480 0.2507 0.2085 0.2595 0.2175 0.1303 0.1716 0.2637 0.0813 0.0081 Ar -0.0352 -0.0352 -0.0045 -0.0227 Lowlands Lowlands

d an l 0.1156 0.1156 0.2085 0.0570 0.2444 0.2865 0.2237 0.0317 0.0216 0.0695 0.2722 0.3355 0.2053 0.0145 0.0339 0.0989 0.0169 0.1280 0.0217 0.0092 0.0171 0.0478 Flats Flats -0.0137 -0.0137 -0.0026 -0.0037 -0.0221 -0.0224 Low

67 67 67 50 50 40 33 33 33 33 33 90 80 80 70 60 50 50 40 40 40 40 33 50 100 100 100 100 % Occ.

*

sp. (annual) sp.

sp. Bromus tectorum Modal Species Species Modal Chamaesyce albomarginata albomarginata Chamaesyce cryptandrusSporobolus Tragopogon dubius cicutarium Erodium serriola Lactuca Astragalus nuttallianus purpurea Aristida dracunculus Artemisia canescens Atriplex pallidum Lycium grossulariifolia Sphaeralcea ramosissima Coleogyne Gilia inconspicua Bromus rubens microcephala Gutierrezia Festuca octoflora Hilaria Cryptantha verna Draba palmerianum Eriogonum Calochortus flexuosus pinnata Descurainia Ephedra nevadensis Stipa hymenoides pallida Oenothera 4. Modal species of vegetation types described by Harper et al. (2001) for Zion National Park, Utah (rows), and their affinitie 4. Modal species of vegetation types described by Harper et al. (2001) for Zion National Park, PPENDIX A Abandoned Fields Vegetation Type Vegetation Type Fields Abandoned Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Abandoned Fields Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Blackbrush Appendix 3). Modal species are prevalent assigned to the vegetation type where their percent occurrence is highest (Cur values (in units of the phi coefficient) between a species and members coalition group. Species are listed in descending o et al. (2003). Green shading by cell value is scaled separately for each column. Nomenclature follows Welsh 76 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 0.0411 0.0411 0.0426 0.1188 0.0487 bank bank -0.0200 -0.0200 -0.0086 -0.0314 -0.0250 -0.0170 -0.0200 -0.0149 -0.1076 -0.1249 -0.0931 -0.0597 -0.0640 -0.0533 -0.0253 -0.0527 -0.0318 -0.0535 -0.0427 -0.0458 -0.1079 -0.0371 -0.0229 -0.0229 -0.0181 -0.0289 Stream- Stream- ce ce i 0.1662 0.1662 0.0517 0.0676 0.0241 0.0364 0.0430 0.0698 -0.0710 -0.0710 -0.0962 -0.0247 -0.0435 -0.0360 -0.0393 -0.0297 -0.0170 -0.1273 -0.0817 -0.0301 -0.0790 -0.0481 -0.0623 -0.0287 -0.0534 -0.0203 -0.0880 -0.0261 -0.0261 -0.0280 -0.0330 Crev Canyon Canyon

h g i 0.0552 0.0552 0.0187 0.0394 0.1026 H -0.0021 -0.0021 -0.1043 -0.0206 -0.0711 -0.0466 -0.0291 -0.0220 -0.1258 -0.0446 -0.0499 -0.0808 -0.0540 -0.0617 -0.0315 -0.0066 -0.0540 -0.0512 -0.0592 -0.0449 -0.0893 -0.0463 -0.0339 -0.0339 -0.0329 -0.0428 Plateau Plateau 0.0173 0.0173 0.0318 0.1598 0.0240 0.1513 0.0374 0.0302 0.0369 0.0278 -0.0307 -0.0307 -0.1114 -0.0274 -0.0726 -0.0710 -0.0420 -0.0263 -0.0930 -0.0145 -0.0210 -0.0256 -0.0425 -0.0522 -0.0770 -0.0609 -0.0537 -0.0550 -0.0550 -0.0593 -0.0660 Upper Upper Mesa Top Top Mesa 0.0226 0.0226 0.1990 0.0472 0.2160 0.0734 0.0074 0.0100 0.0113 -0.0650 -0.0650 -0.1590 -0.0536 -0.0941 -0.0914 -0.0115 -0.0589 -0.0431 -0.1054 -0.0038 -0.0414 -0.0231 -0.0254 -0.0605 -0.0952 -0.0401 -0.0459 -0.0626 -0.0626 -0.0675 -0.0747 Lower Lower Mesa Top Top Mesa 0.0141 0.0141 0.0178 0.2217 0.0704 0.0148 0.3022 0.2407 0.1966 0.2903 0.0551 0.2097 -0.0941 -0.0941 -0.0984 -0.0282 -0.0495 -0.0587 -0.0339 -0.0255 -0.1486 -0.0081 -0.1071 -0.0519 -0.0797 -0.0076 -0.0142 -0.0417 -0.0417 -0.0388 -0.0532

d an l 0.0087 0.0087 0.0133 0.1202 0.0060 0.0340 0.0173 0.0150 0.0211 0.1001 0.1601 0.1432 0.0973 0.0948 0.1880 0.1933 0.0831 0.2100 0.1172 -0.0696 -0.0696 -0.0568 -0.0308 -0.0339 -0.0462 -0.0616 -0.0189 -0.0395 -0.0395 -0.0542 -0.0679 Sands Slickrock Slickrock Sands Up y k 0.0140 0.0140 0.0670 0.0099 0.0303 0.0203 0.0187 0.1868 0.1661 0.2379 0.1964 0.0969 0.1502 0.0990 0.2064 0.0883 0.0028 0.0206 0.1969 0.0846 0.0095 0.0095 -0.0212 -0.0212 -0.0321 -0.0358 -0.0008 -0.0408 -0.0185 -0.0240 -0.0240 -0.0107 Roc Slopes Slopes

id 0.0237 0.0237 0.2817 0.0923 0.0136 0.1752 0.0449 0.0666 0.1706 0.0295 0.1617 0.0388 0.0284 0.0051 0.2058 0.2058 0.1065 0.2031 Ar -0.0041 -0.0041 -0.0968 -0.0799 -0.0387 -0.0898 -0.0340 -0.0651 -0.0732 -0.0513 -0.0529 -0.0544 -0.0310 Lowlands Lowlands

d an l 0.0211 0.0211 0.0989 0.0162 0.0561 0.1030 0.0226 0.0251 0.0487 0.0487 0.0240 Flats Flats -0.0081 -0.0081 -0.0097 -0.0900 -0.0953 -0.0639 -0.0363 -0.0361 -0.0219 -0.0291 -0.0775 -0.0036 -0.0498 -0.0741 -0.0544 -0.0433 -0.0392 -0.0871 -0.0564 -0.0062 -0.0062 Low

71 57 43 29 29 29 29 82 65 57 53 34 31 29 33 32 32 32 84 64 64 60 56 48 30 30 30 100 100 % Occ.

1 tenuifolia Artemisia tridentata tridentata Artemisia sarothrae Gutierrezia elymoides Elymus nauseosus Chrysothamnus sparsiflorum Eriastrum divergens Erigeron Opuntia phaeacantha setosa Pectocarya osteosperma Juniperus utahensis Amelanchier Arabis holboellii Pinus monophylla edulis Pinus Streptanthus cordatus Erigeron sionis racemosum Eriogonum asperum Erysimum canescens Machaeranthera heterophylla Phacelia intricatus Cercocarpus fendleri Arenaria villosa Chrysopsis scabrida Castilleja macrorhiza Opuntia Stephanomeria Modal Species Species Modal Eriastrum eremicum eremicum Eriastrum Poa bigelovii fremontii Psorothamnus 4. Continued. PPENDIX A Sagebrush Sagebrush Sagebrush Sagebrush Sagebrush Sagebrush Sagebrush Sagebrush Sagebrush Juniper-Pinyon Juniper-Pinyon Juniper-Pinyon Juniper-Pinyon Juniper-Pinyon Juniper-Pinyon Juniper-Pinyon Mountain Brush Mountain Brush Mountain Brush Mountain Brush Rock Crevice Rock Crevice Rock Crevice Rock Crevice Rock Crevice Rock Crevice Vegetation Type Vegetation Type Blackbrush Blackbrush Blackbrush Blackbrush 2015] PLANT COALITION GROUPS OF ZION NP 77 0.0410 0.0410 0.0212 0.0130 0.0429 0.0000 bank bank -0.0456 -0.0456 -0.0586 -0.0469 -0.0438 -0.0524 -0.0316 -0.0184 -0.0322 -0.0837 -0.0949 -0.0606 -0.0675 -0.0010 -0.0475 -0.0588 -0.0505 -0.0521 -0.0480 -0.0386 -0.0554 -0.0198 -0.0142 -0.0515 Stream- Stream- ce ce i 0.0411 0.0411 0.0259 0.1111 0.0285 0.0666 0.1121 0.0042 0.2236 0.2559 0.0347 0.2614 0.1409 0.1310 -0.0342 -0.0342 -0.0409 -0.0403 -0.0597 -0.0072 -0.0837 -0.0054 -0.0503 -0.0403 -0.0563 -0.0037 -0.0470 -0.0344 -0.0018 -0.0055 Crev Canyon Canyon

h g i H 0.0460 0.0460 0.1711 0.1418 0.0248 0.1027 0.1859 0.2005 0.0309 0.2497 0.1139 0.0752 -0.0542 -0.0542 -0.0622 -0.0683 -0.0374 -0.0805 -0.0192 -0.0050 -0.0417 -0.0959 -0.0288 -0.0512 -0.0297 -0.0401 -0.0146 -0.0629 -0.0115 -0.0509 Plateau Plateau 0.0032 0.0032 0.0717 0.0891 0.0301 0.0478 0.0433 0.0855 0.1832 0.0821 0.1804 0.1310 0.1553 0.0229 0.1700 0.0042 0.1074 0.0003 0.0067 0.0324 0.0969 -0.0335 -0.0335 -0.0637 -0.0125 -0.0825 -0.0131 -0.0091 -0.0069 -0.0226 Upper Upper Mesa Top Top Mesa 0.0168 0.0168 0.0101 0.0611 0.0577 0.0893 0.0662 0.0491 0.0050 0.1318 0.0257 0.1792 0.2096 0.2114 0.0603 0.1381 0.0553 0.0806 0.0312 0.0802 0.0255 0.1614 -0.0416 -0.0416 -0.0008 -0.0691 -0.0007 -0.0980 -0.0052 -0.0337 Lower Lower Mesa Top Top Mesa 0.0846 0.0846 0.1140 0.0972 0.2021 0.2338 0.1297 0.1993 0.1186 0.0767 0.1136 0.0721 0.0311 0.1502 0.0282 0.0550 0.0422 0.0706 0.1222 0.0865 0.1328 0.0587 0.2068 0.0046 -0.0280 -0.0280 -0.0355 -0.0478 -0.0499 -0.0046

d an l 0.2316 0.2316 0.2527 0.0887 0.1052 0.0456 0.1198 0.1735 0.1283 0.2147 0.1919 0.1886 0.1548 0.0959 0.0990 0.1314 0.1379 0.0354 0.0118 0.0505 0.0971 0.0711 Sands Slickrock Slickrock Sands -0.0137 -0.0137 -0.0559 -0.0492 -0.0183 -0.0652 -0.0489 -0.0508 Up y k 0.0773 0.0773 0.1114 0.1252 0.0790 0.2131 0.0496 0.2132 0.1132 0.1615 0.1251 0.0241 0.1271 0.0777 0.1162 0.0124 0.1804 Roc -0.0209 -0.0209 -0.0675 -0.0321 -0.0259 -0.0118 -0.0482 -0.0770 -0.0154 -0.0415 -0.0621 -0.0451 -0.0511 Slopes Slopes

id Ar 0.0704 0.0704 0.1136 -0.0162 -0.0162 -0.0719 -0.0565 -0.0719 -0.0415 -0.1236 -0.0620 -0.0593 -0.1087 -0.0520 -0.0693 -0.0642 -0.0607 -0.0696 -0.0505 -0.2123 -0.1407 -0.1431 -0.0893 -0.1199 -0.0849 -0.0951 -0.0988 -0.0823 -0.0438 -0.0947 Lowlands Lowlands

d an l Flats Flats -0.0532 -0.0532 -0.0625 -0.0351 -0.0593 -0.0617 -0.0448 -0.0179 -0.0574 -0.1391 -0.1013 -0.0802 -0.0767 -0.0015 -0.0398 -0.0804 -0.0686 -0.0660 -0.0835 -0.0573 -0.0335 -0.0226 -0.0154 -0.0476 -0.0117 -0.0556 -0.0516 -0.0620 -0.0231 Low

40 36 36 36 32 28 28 97 85 61 52 39 33 30 24 24 24 86 71 57 52 52 48 48 43 38 38 38 % Occ. Modal Species Species Modal Penstemon laevis laevis Penstemon Chaenactis douglasii Cryptantha confertiflora humilis Penstemon Quercus turbinella canaani Erigeron rostriflorus Penstemon Pinus ponderosa fendleriana Poa patula Arctostaphylos Senecio multilobatus austromontana Phlox Polygonum douglasii tridentata Purshia macradenia Arenaria Comandra umbellata angustissima Yucca gambelii Quercus myrsinites Pachystima rossii Carex concolor Abies oreophilus Symphoricarpos menziesii Pseudotsuga americana Vicia Acer grandidentatum Draba asprella Haplopappus scopulorum Mahonia repens 4. Continued. PPENDIX A Vegetation Type Vegetation Type Rock Crevice Rock Crevice Rock Crevice Rock Crevice Rock Crevice Rock Crevice Rock Crevice Pine Ponderosa Pine Ponderosa Pine Ponderosa Pine Ponderosa Pine Ponderosa Pine Ponderosa Pine Ponderosa Ponderosa Pine (cont.) Pine Ponderosa Pine Ponderosa Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas 78 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 ------0.0442 0.0442 0.2513 0.1026 0.3353 0.0380 0.0380 0.0109 0.1134 0.0817 0.1156 0.0462 0.0088 0.2268 0.0911 0.0540 bank bank -0.0222 -0.0222 -0.0520 -0.0007 -0.0025 -0.0085 -0.0085 -0.0070 -0.0025 -0.0134 -0.0560 -0.0182 -0.0060 Stream- Stream- ce ce i ------0.1183 0.1183 0.3163 0.0962 0.0071 0.1947 0.1947 0.2533 0.0534 0.0134 0.1540 0.0179 0.1633 0.0179 0.0585 0.0387 0.2110 0.1160 0.3612 0.1476 0.1274 0.0811 0.0379 0.0093 0.0954 0.1447 -0.0092 -0.0092 -0.0588 Crev Canyon Canyon

h g i 0.0007 0.0007 0.0499 0.0195 0.0476 0.0476 0.3314 0.1754 0.0132 0.0224 0.0476 0.0135 0.0630 0.0889 0.0284 0.0068 0.0262 0.0635 H -0.0126 -0.0126 -0.0027 -0.0027 -0.0571 -0.0127 -0.0127 -0.0163 -0.0025 -0.0713 -0.0130 -0.0274 Plateau Plateau 0.0020 0.0020 0.0497 0.0409 0.1036 0.0148 -0.0252 -0.0252 -0.0579 -0.0201 -0.0647 -0.0308 -0.0308 -0.0019 -0.0308 -0.0217 -0.0375 -0.0217 -0.0232 -0.0086 -0.0030 -0.0173 -0.0365 -0.0681 -0.0243 -0.0274 -0.0142 -0.0217 -0.0078 Upper Upper Mesa Top Top Mesa

------0.0824 0.0824 0.0382 0.0271 0.0539 0.0152 0.0054 0.0614 0.0020 0.0102 0.0028 0.0195 -0.0282 -0.0282 -0.0447 -0.0376 -0.0734 -0.0046 -0.0046 -0.0352 -0.0248 -0.0372 -0.0248 -0.0250 -0.0123 -0.0054 -0.0700 -0.0207 -0.0248 Lower Lower Mesa Top Top Mesa

------0.0736 0.0736 0.0346 0.3078 0.3078 0.0183 0.1744 0.0919 0.0270 0.1492 0.1890 0.3143 0.2838 0.1052 0.0238 -0.0231 -0.0231 -0.0407 -0.0308 -0.0308 -0.0283 -0.0028 -0.0163 -0.0144 -0.0163 -0.0108 -0.0091 -0.0274 -0.0215 -0.0163

d an l 0.0261 0.0261 0.0077 0.0077 0.0046 0.0464 0.0268 0.0809 0.0434 -0.0340 -0.0340 -0.0678 -0.0613 -0.0204 -0.0204 -0.0292 -0.0712 -0.0170 -0.0340 -0.0240 -0.0507 -0.0240 -0.0269 -0.0340 -0.0189 -0.0389 -0.0391 -0.0334 -0.0240 -0.0057 Sands Slickrock Slickrock Sands Up y k 0.1161 0.1161 0.2408 0.2408 0.0115 0.2246 0.0075 -0.0466 -0.0466 -0.0929 -0.0857 -0.0438 -0.0438 -0.0461 -0.1064 -0.0461 -0.0466 -0.0368 -0.0609 -0.0368 -0.0509 -0.0469 -0.0414 -0.0670 -0.0889 -0.0582 -0.0526 -0.0640 -0.0329 -0.0021 Roc Slopes Slopes

id 0.0007 0.0007 0.0431 0.0431 0.1342 Ar -0.0260 -0.0260 -0.0711 -0.0336 -0.0921 -0.0921 -0.0709 -0.0822 -0.0775 -0.0290 -0.0204 -0.0449 -0.0204 -0.0355 -0.0290 -0.0290 -0.0144 -0.0709 -0.1190 -0.0612 -0.0610 -0.0355 -0.0183 -0.0538 -0.0276 Lowlands Lowlands

d an l 0.0312 0.0312 0.0048 0.0048 0.0085 0.0236 Flats Flats -0.0154 -0.0154 -0.0226 -0.0089 -0.0356 -0.0356 -0.0588 -0.0292 -0.0178 -0.0154 -0.0109 -0.0269 -0.0109 -0.0189 -0.0154 -0.0154 -0.0269 -0.0081 -0.0341 -0.0385 -0.0174 -0.0189 -0.0109 -0.0189 Low

72 63 42 42 28 -- -- 28 ------56 -- -- 56 ------33 29 29 29 24 89 79 72 50 50 44 40 40 40 40 39 39 33 33 28 26 20 % Occ.

2 2 Modal Species Species Modal Mimulus cardinalis cardinalis Mimulus Acer negundo ludoviciana Artemisia Bromus diandrus Holodiscus dumosus Holodiscus Fraxinus anomala rubescens Heuchera jamesiana Stellaria scopulorum Juniperus capillus-veneris Adiantum scopulorum Calamagrostis welshii Aster cardinalis Lobelia chrysantha Aquilegia formosa Aquilegia pulchellum Dodecatheon androsaemifolium Apocynum glaucodes Aster Eupatorium herbaceum velutina Solidago utahensis Erigeron caespitosum Petrophytum thurberi Muhlenbergia Smilacina stellata stolonifera Agrostis curatorum Carex arizonicum Cirsium Antennaria rosulata 4. Continued. PPENDIX A Vegetation Type Vegetation Type Riparian Riparian Riparian Riparian Douglas Fir-White Fir Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Fir Fir-White Douglas Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging Gardens Hanging 2015] PLANT COALITION GROUPS OF ZION NP 79 t bank bank 0.0995 0.0995 0.0200 0.1862 0.3936 0.3744 0.0691 0.2027 0.4269 0.0474 -0.0039 -0.0039 Stream- Stream- ce ce i 0.2602 0.2602 0.1054 0.0263 0.0149 0.1603 0.1956 0.3184 0.0391 0.0266 0.1166 Crev Canyon Canyon

h g i H 0.0473 0.0473 0.0085 0.0008 0.0372 0.0217 0.0764 0.2466 0.0066 0.0083 -0.0130 -0.0130 Plateau Plateau Upper Upper -0.0460 -0.0460 -0.0629 -0.0496 -0.0487 -0.0537 -0.0293 -0.0366 -0.0258 -0.0489 -0.0249 Mesa Top Top Mesa Lower Lower -0.0176 -0.0176 -0.0538 -0.0546 -0.0613 -0.0501 -0.0038 -0.0160 -0.0455 -0.0559 -0.0327 Mesa Top Top Mesa 0.0757 0.0757 0.0257 0.2515 0.0833 0.0139 -0.0313 -0.0313 -0.0362 -0.0190 -0.0409 -0.0322

d an l 0.0456 0.0456 0.0005 Sands Slickrock Slickrock Sands -0.0486 -0.0486 -0.0478 -0.0534 -0.0464 -0.0311 -0.0276 -0.0476 -0.0046 Up y k 0.0363 0.0363 Roc Slopes Slopes -0.0665 -0.0665 -0.0308 -0.0638 -0.0756 -0.0812 -0.0283 -0.0547 -0.0991 -0.0696

id Ar -0.0670 -0.0670 -0.0684 -0.0349 -0.0177 -0.0422 -0.0558 -0.0505 -0.0598 -0.0253 -0.0363 Lowlands Lowlands

d an l Flats Flats 0.0293 0.0293 0.0242 0.0155 -0.0242 -0.0242 -0.0169 -0.0139 -0.0160 -0.0045 -0.0245 -0.0291 Low

37 37 32 32 26 26 21 21 21 21 % Occ. were present in supplemental hanging garden data used by Harper et al. (2001) but not used but in et al. (2001) Harper present in garden by hanging data were used supplemental were present in supplemental hanging garden data used by Harper et al. (2001) but not coalition group analysis. Modal Species Species Modal Bromus ciliatus Senecio spartioides Fraxinus velutina fremontii Populus hyemale Equisetum Zauschneria latifolia fragilis Cystopteris pratensis Poa chinensis Tamarix Thelypodium laxiflorum listed in Harper et al. (2001) were lumped listedthe in were in Harper et al. (2001) uncer duecoalition group identification to analysis Lobelia cardinalis Lobelia listed in Harper et al. (2001) were lumped the coalition group analysis due to identification uncertainties. and and Lobelia cardinalis Gutierrezia and 4. Continued. Gutierrezia PPENDIX A Vegetation Type Vegetation Type Riparian Riparian Riparian Riparian Riparian Riparian Riparian Riparian Riparian Riparian The species of species of The curatorumCarex The species of Carex curatorum

1 2 1 2 80 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 0.0039 0.0039 bank -0.0214 -0.0214 -0.0254 -0.0261 -0.0407 -0.0268 -0.0565 -0.0622 -0.0427 -0.0286 -0.0477 -0.0663 -0.0521 -0.0431 -0.0450 -0.0758 -0.0528 -0.0307 -0.0308 -0.0416 -0.0190 -0.0228 -0.0399 -0.0196 -0.0356 Stream- 0.0121 0.0121 0.0486 0.0246 -0.0105 -0.0105 -0.0217 -0.0170 -0.0347 -0.0461 -0.0143 -0.0178 -0.0215 -0.0351 -0.0201 -0.0160 -0.0318 -0.0561 -0.0391 -0.0272 -0.0223 -0.0314 -0.0227 -0.0024 -0.0169 -0.0183 -0.0382 Crevice Crevice Canyon Canyon 0.0427 0.0427 0.0639 0.0242 0.0357 0.0195 0.0420 0.0114 0.0101 0.0116 0.0164 0.0113 0.0083 High -0.0280 -0.0280 -0.0317 -0.0107 -0.0313 -0.0080 -0.0049 -0.0037 -0.0393 -0.0386 -0.0117 -0.0183 -0.0125 -0.0194 Plateau 0.0639 0.0639 0.0961 0.0651 0.0747 0.0847 0.0811 0.0628 0.0808 0.0632 0.0460 0.0761 0.0780 0.0665 0.0651 0.0580 0.0598 0.0410 0.0523 0.0182 0.0230 0.0041 0.0486 0.0363 0.0166 0.0143 Upper Mesa Top . (2004), placed within an alternative classi- 0.0531 0.0531 0.1158 0.0921 0.0564 0.0817 0.0902 0.0845 0.1039 0.0699 0.0363 0.0814 0.0983 0.0718 0.0768 0.0340 0.0742 0.0577 0.0264 0.0062 0.0241 0.0383 0.0085 -0.0001 -0.0001 -0.0099 -0.0101 Lower Lower Mesa Top cell value is scaled separately for each column. ts of the phi coefficient. The values are calculated for each 0.0331 0.0331 0.0408 0.0317 0.0106 0.0052 0.0079 0.0263 0.0309 0.0215 0.0109 0.0219 0.0304 0.0425 -0.0002 -0.0002 -0.0312 -0.0053 -0.0176 -0.0058 -0.0032 -0.0391 -0.0123 -0.0221 -0.0022 -0.0108 -0.0041 0.0478 0.0478 0.0176 0.0303 0.0294 0.0305 0.0301 0.0229 0.0140 0.0343 0.0486 0.0627 0.0715 0.0660 0.0361 0.0158 0.0272 0.0313 0.0965 0.1264 0.0956 0.0686 0.0912 0.0458 0.0474 -0.0127 -0.0127 Sands Slickrock Upland 0.0136 0.0136 0.0015 0.0225 0.0250 0.0439 0.0768 0.0898 0.0604 0.0641 0.0714 0.0953 0.0729 0.0357 0.0468 0.1151 0.0965 0.0392 0.0485 0.0579 0.0227 0.0558 0.0560 0.0503 0.0704 -0.0104 -0.0104 Rocky Slopes 0.0065 0.0065 0.0230 -0.0601 -0.0601 -0.0862 -0.0717 -0.0548 -0.0587 -0.0711 -0.0268 -0.0230 -0.0304 -0.0211 -0.0447 -0.0407 -0.0501 -0.0507 -0.0101 -0.0051 -0.0072 -0.0197 -0.0387 -0.0280 -0.0376 -0.0437 -0.0193 Arid

Lowlands 0.0223 0.0223 0.0025 Flats -0.0295 -0.0295 -0.0442 -0.0425 -0.0296 -0.0329 -0.0364 -0.0374 -0.0360 -0.0297 -0.0449 -0.0542 -0.0488 -0.0408 -0.0180 -0.0517 -0.0407 -0.0229 -0.0460 -0.0199 -0.0330 -0.0444 -0.0139 -0.0031

Lowland Woodland / Shrubland / Herbaceous / Sparse / Herbaceous / Shrubland Woodland - (Streamside Wild Rye) Shrubland Rye) Wild - (Streamside ead Herbaceous Vegetationead Herbaceous

5. Associations of the U.S. National Vegetation Classification System at Zion National Park, Utah, as recognized by Cogan et al Classification System at Zion National Park, National Vegetation 5. Associations of the U.S. PPENDIX A XXx. Alternative Classification HierarchyClassification Alternative XXx. Association NVC VEGETATION ZION A. XERIC Slopes / Rocky / Slickrock Sands Upland Top / Mesa A1. A1a. Mesa Top - / Forest / Woodland Shrubland Woodland Oak / Gambel Pine Ponderosa Forest Manzanita / Greenleaf Fir White Shrubland Manzanita Mexican Woodland Sagebrush / Black Pine Ponderosa Woodland Mountain-mahogany / Curl-leaf Pinyon Two-needle Wheatgrass / Western Sagebrush Big Basin Sparse / / Shrubland Woodland - Slopes Rocky Top / Mesa A1b. Vegetation Mountain-mahogany Sparse Pavement Rock / Mountain-mahogany species - Woodland Juniper Pinyon Two-needle - Sedge) Shrubland Oak Mountain-mahogany(Geyer's Gambel / Shrubland Serviceberry / Utah Oak Gambel Woodland Oak / Gambel species - Juniper Pinyon Two-needle Woodland Manzanita / Greenleaf - Juniper Utah Pinyon Two-needle Shrubland - (Utah Oak - Serviceberry) Gambel Manzanita Greenleaf Manzanita Woodland / Greenleaf Pine Ponderosa Dwarf-shrubland / Muttongrass Sagebrush Black Woodland - / Mountain-mahogany Oak Gambel - Juniper Utah Pinyon Singleleaf Woodland Cliff-rose / Stansbury - Juniper Utah Pinyon Two-needle Sagebrush Big / Needle-and-ThreadMountain Shrubland - Slopes Rocky / / Slickrock Sands Upland A1c. Blue Grama - Needle-and-Thr - Mountain Big Shrubland Manzanita Sagebrush Greenleaf Shrubland Grama / Blue Sagebrush Big Shrubland / Muttongrass Oak Gambel Shrubland Manzanita Greenleaf Shrubland Sagebrush / Big Oak Gambel Sagebrush / Needle-and-Thread Dwarf-shrubland Black fication hierarchy based on affinities to coalition groups (see Fig. 7 in paper). Affinity values (right 10 columns) are in uni 7 in paper). Affinity fication hierarchy based on affinities to coalition groups (see Fig. species in relation to each coalition group (see Appendix 3) and then averaged across plots associations. Green shading by 2015] PLANT COALITION GROUPS OF ZION NP 81 0.0469 0.0469 0.0424 0.0710 0.0275 bank -0.0470 -0.0470 -0.0409 -0.0622 -0.0436 -0.0362 -0.0362 -0.0148 -0.0083 -0.0127 -0.0115 -0.0251 -0.0139 -0.0165 -0.0327 -0.0432 -0.0289 -0.0204 -0.0471 -0.0563 -0.0389 -0.0004 -0.0074 Stream- -0.0479 -0.0479 -0.0271 -0.0422 -0.0344 -0.0082 -0.0082 -0.0156 -0.0027 -0.0374 -0.0181 -0.0430 -0.0461 -0.0524 -0.0405 -0.0596 -0.0605 -0.0653 -0.0478 -0.0540 -0.0696 -0.0844 -0.0604 -0.0421 -0.0477 -0.0204 -0.0229 Crevice Crevice Canyon Canyon 0.0378 0.0378 0.0349 High -0.0387 -0.0387 -0.0354 -0.0432 -0.0448 -0.0417 -0.0417 -0.0167 -0.0563 -0.0096 -0.0556 -0.0460 -0.0620 -0.0517 -0.0669 -0.0695 -0.0675 -0.0555 -0.0542 -0.0647 -0.0754 -0.0604 -0.0511 -0.0430 -0.0464 Plateau 0.0186 0.0186 0.0204 0.0246 0.0184 0.0350 0.0350 -0.0008 -0.0008 -0.0024 -0.0820 -0.0485 -0.0827 -0.0631 -0.0648 -0.0684 -0.0797 -0.0595 -0.0701 -0.0491 -0.0451 -0.0263 -0.0336 -0.0398 -0.0525 -0.0445 -0.0713 -0.0258 Upper Mesa Top 0.0112 0.0112 0.0123 0.0244 0.0221 0.0052 0.0052 -0.0043 -0.0043 -0.0134 -0.0995 -0.0718 -0.0967 -0.0758 -0.0783 -0.0852 -0.1016 -0.0745 -0.0861 -0.0599 -0.0661 -0.0479 -0.0558 -0.0555 -0.0690 -0.0660 -0.0746 -0.0465 Lower Lower Mesa Top 0.0381 0.0381 0.0418 0.0180 0.1010 0.1010 0.0557 -0.0180 -0.0180 -0.0086 -0.0086 -0.0301 -0.0465 -0.0608 -0.0576 -0.0509 -0.0509 -0.0600 -0.0600 -0.0706 -0.0422 -0.0471 -0.0693 -0.0698 -0.0564 -0.0316 -0.0469 -0.0368 -0.0382 0.0806 0.0806 0.0791 0.0913 0.0582 0.0899 0.0899 0.1097 0.0385 0.0175 0.0031 0.0103 0.0151 0.0545 0.0057 -0.0194 -0.0194 -0.0627 -0.0267 -0.0258 -0.0204 -0.0409 -0.0290 -0.0488 -0.0180 -0.0068 -0.0079 -0.0170 Sands Slickrock Upland 0.0675 0.0675 0.0941 0.0903 0.1013 0.0572 0.0572 0.0481 0.0041 0.0181 0.0137 0.0064 0.0241 0.0207 0.0421 0.0466 0.0586 0.0527 0.0503 0.0856 0.0947 0.0825 0.0369 0.0419 0.0273 -0.0038 -0.0038 -0.0073 Rocky Slopes 0.0012 0.0012 0.0053 0.0819 0.0819 0.0821 0.1541 0.1107 0.1379 0.1486 0.1470 0.1692 0.2037 0.1338 0.1171 0.1322 0.1489 0.1264 0.0861 0.0781 0.0450 0.0253 -0.0118 -0.0118 -0.0157 -0.0225 -0.0225 -0.0153 -0.0171 Arid Lowlands 0.0212 0.0212 0.0686 0.0752 0.1295 0.0954 0.0864 0.0961 0.0877 0.0591 0.0477 0.0203 0.0557 0.0036 0.0212 0.0131 0.0276 0.0364 0.0100 0.0508 Flats -0.0224 -0.0224 -0.0389 -0.0464 -0.0395 -0.0428 -0.0428 -0.0305

Lowland aceous Vegetation atural Shrub Herb Bluegrass Semi-n sin Herbaceous Vegetationsin Flooded Shrubland en Sparse Vegetation en

eat Basin Herbaceous Vegetation Herbaceous Basin eat Basalt Shrubland Basalt 5. Continued. PPENDIX A XXx. Alternative Classification HierarchyClassification Alternative XXx. Association NVC Woodland Sagebrush / Big species - Juniper Pinyon Two-needle Vegetation Sparse Slickrock Pine Ponderosa Shrubland Manzanita - Cliff-rose Greenleaf Stansbury Woodland Mountain-mahogany / Littleleaf - Juniper Utah Pinyon Two-needle Littleleaf Mountain-mahogany Slickrock Sparse Vegetation Mountain-mahogany Sparse Slickrock Littleleaf (Sandhill Muhly, MountainMuhly) - Hairy Goldenaster Herbaceous Vegetation A1 outlier Needle-and-Thread Great Ba Sands / Upland Slopes / Rocky A2. Lowlands / Sparse / Herbaceous / Shrubland Woodland Flats - / Lowland Lowlands Arid A2a. Sand Dropseed Gr Arrow-weed Seasonally Nevada Joint-fir / Lich Shrubland - Sagebrush Big Saltbush Fourwing Shrubland Saltbush Fourwing Nevada Joint-fir Vegetation Herbaceous Galleta James' Sparse / / Shrubland Woodland - Lowlands Arid A2b. Blackbrush Shrubland Shrubland / Galleta James' Blackbrush Vegetation Sparse Badlands Buckwheat Wild Crispleaf Dwarf-shrubland / James' Galleta species) - (Prickly-pear Snakeweed Woodland / Blackbrush - Juniper Utah Pinyon Singleleaf Woodland / Big Sagebrush Juniper Utah Woodland Sagebrush / Big - Juniper Utah Pinyon Singleleaf Shrubland Plateau Colorado Sagebrush Sand Shrubland Semi-natural / Cheatgrass Rabbitbrush) - Sagebrush Big (Rubber A2 outliers Coyote Willow / Barren Shrubland Green Rabbitbrush / Kentucky 82 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 0.0274 0.0274 0.0213 0.1332 0.0000 0.0218 0.0368 0.0664 0.0634 0.0585 0.0341 0.0174 0.0400 0.0457 0.0072 0.0072 bank -0.0068 -0.0068 -0.0095 -0.0201 -0.0201 -0.0212 -0.0219 -0.0373 -0.0474 -0.0531 -0.0459 -0.0274 -0.0141 Stream- on y 0.0703 0.0703 0.1288 0.0569 0.0551 0.0897 0.0855 0.0378 0.0337 0.1435 0.1910 0.0380 0.0064 0.0021 0.0457 0.0609 -0.0184 -0.0184 -0.0537 -0.0328 -0.0248 -0.0490 -0.0480 -0.0598 -0.0543 -0.0253 -0.0067 Crevice Crevice Can 0.0782 0.0782 0.0785 0.0198 0.0520 0.1057 0.0487 0.0707 0.0901 0.0869 0.0481 0.0282 0.1086 0.0717 0.1285 0.1860 High -0.0293 -0.0293 -0.0707 -0.0380 -0.0082 -0.0569 -0.0549 -0.0555 -0.0356 -0.0287 -0.0109 Plateau

p 0.0282 0.0282 0.0347 0.0536 0.0788 0.0436 0.0077 0.0337 0.0067 0.0170 0.0234 0.0624 0.0221 0.0391 0.0052 0.0052 0.0019 -0.0145 -0.0145 -0.0206 -0.0413 -0.0413 -0.0495 -0.0125 -0.0259 -0.0156 -0.0002 -0.0012 -0.0074 Upper Mesa To

p 0.0316 0.0316 0.0524 0.0635 0.0957 0.0662 0.0049 0.0145 0.0435 0.0133 0.0173 0.0100 0.0256 -0.0080 -0.0080 -0.0146 -0.0113 -0.0492 -0.0492 -0.0668 -0.0324 -0.0173 -0.0365 -0.0223 -0.0129 -0.0180 -0.0055 -0.0076 Lower Lower Mesa To 0.0228 0.0228 0.0521 0.0144 0.0298 0.0440 0.0477 0.0498 0.0429 0.0506 0.0594 0.0073 0.0024 0.0060 0.0060 0.0032 0.0094 -0.0025 -0.0025 -0.0124 -0.0049 -0.0269 -0.0269 -0.0245 -0.0195 -0.0413 -0.0448 -0.0017 -0.0133 0.0090 0.0090 0.0063 0.0082 0.0117 0.0398 0.0371 0.0206 0.0634 0.0634 0.0697 0.0993 0.0238 0.0272 0.0204 0.0154 0.0332 0.0149 -0.0019 -0.0019 -0.0006 -0.0152 -0.0182 -0.0045 -0.0175 -0.0046 -0.0256 -0.0188 -0.0188 Sands Slickrock Upland es p 0.0114 0.0114 0.0161 0.0252 0.0318 0.0318 0.0925 0.0589 0.0215 0.1121 0.1224 0.1077 0.0775 0.0948 0.0902 -0.0086 -0.0086 -0.0156 -0.0341 -0.0198 -0.0254 -0.0214 -0.0443 -0.0487 -0.0411 -0.0070 -0.0404 -0.0505 Rocky Slo 0.0194 0.0194 0.0489 0.0029 0.0195 0.0753 0.0722 0.0704 0.0719 0.0281 0.0288 -0.0669 -0.0669 -0.0767 -0.0347 -0.0557 -0.0982 -0.0697 -0.0532 -0.0553 -0.0499 -0.0773 -0.0325 -0.0225 -0.0306 -0.0447 -0.0594 Arid Lowlands 0.0581 0.0581 0.0931 0.0062 0.0115 0.0115 0.0036 0.0176 Flats -0.0266 -0.0266 -0.0367 -0.0211 -0.0345 -0.0408 -0.0461 -0.0297 -0.0197 -0.0029 -0.0368 -0.0309 -0.0067 -0.0231 -0.0231 -0.0164 -0.0236 -0.0152 -0.0010 -0.0223 -0.0310 Lowland

Bluegrass Semi-natural Forest Forest Semi-natural Bluegrass wberry / Tall Forbs Forest Forbs / Tall wberry - Gambel Oak Woodland Brickelbush Woodland Semi-natural Seasonally Flooded Herbaceous Alliance

5. Continued. PPENDIX A XXx. Alternative Classification HierarchyClassification Alternative XXx. Association NVC B. MESIC ZION VEGETATION ZION VEGETATION B. MESIC Canyon / Crevice Plateau B1. High / Herbaceous / Woodland - Forest Canyon / Crevice Plateau B1a. High Forest Oak Maple / Gambel Bigtooth Maple Forest / Bigtooth Fir White Vegetation Sedge Herbaceous Beaked Douglas-fir / Mountain Snowberry Forest Forest Oak / Gambel Fir White Forest Oak / Gambel Douglas-fir Woodland Bracken / Northern Pine Ponderosa Woodland Semi-natural / Smooth Brome Pine Ponderosa / Tasselflower Box-elder Maple Forest / Bigtooth Douglas-fir Rocky Mountain Juniper / Herbaceous / Shrubland Woodland / - Forest Plateau B1b. High Kentucky Bluegrass Vegetation Semi-natural Herbaceous Wheatgrass) - Brome Smooth (Western / Kentucky Fir Aspen - White Quaking Sno Aspen / Mountain Quaking A2c. Upland Sands / Rocky Slopes - Shrubland / Sparse Sparse / Shrubland - Slopes / Rocky Sands Upland A2c. Sparse Vegetation Sand Deposit Rabbitbrush Rubber Semi-natural Shrubland / Cheatgrass Rabbitbrush Rubber - Mormon-tea Shrubland Horsebrush Gray Dwarf-shrubland / Bottlebrush Sagebrush Black A2d. Rocky Slopes / - Woodland Shrubland Woodland Live Oak / Turbinella - Juniper Utah Pinyon Singleleaf Woodland Serviceberry) Utah Buffaloberry, / (Roundleaf - Juniper Utah Pinyon Singleleaf Woodland Understory / Sparse - Juniper Utah Pinyon Singleleaf Woodland Sagebrush / Black - Juniper Utah Pinyon Singleleaf Shrubland Colluvial - Serviceberry) (Utah Live Oak Turbinella Ash Woodland Single-leaf 2015] PLANT COALITION GROUPS OF ZION NP 83 0.0600 0.0600 0.0487 0.0214 0.0038 0.0061 0.0104 0.0181 0.1606 0.1540 0.1064 0.1149 0.0505 0.2246 0.2544 0.2354 0.1710 0.1466 0.1520 bank Stream- 0.0604 0.0604 0.0408 0.0096 0.0207 0.0412 0.0659 0.0168 0.0526 0.0315 0.0421 0.0733 0.0480 0.0431 0.0475 0.0219 0.0370 0.0046 0.0069 Crevice Crevice Canyon Canyon 0.1635 0.1635 0.1599 0.1238 0.1088 0.1542 0.1266 0.0651 0.0671 0.0801 0.0448 0.1070 0.0434 0.0225 0.0180 0.0297 High -0.0128 -0.0128 -0.0098 -0.0114 Plateau 0.0497 0.0497 0.0349 0.0253 0.0683 0.0731 0.0589 0.0253 0.0005 -0.0137 -0.0137 -0.0133 -0.0126 -0.0165 -0.0396 -0.0250 -0.0468 -0.0400 -0.0355 -0.0381 Upper Mesa Top 0.0270 0.0270 0.0089 0.0036 0.0521 0.0594 0.0517 0.0201 -0.0189 -0.0189 -0.0267 -0.0014 -0.0084 -0.0242 -0.0408 -0.0315 -0.0569 -0.0414 -0.0474 -0.0471 Lower Lower Mesa Top 0.0001 0.0001 0.0001 0.0165 0.0197 0.0009 -0.0147 -0.0147 -0.0092 -0.0003 -0.0245 -0.0073 -0.0030 -0.0027 -0.0025 -0.0087 -0.0139 -0.0161 -0.0152 -0.0186 0.0102 0.0102 0.0096 0.0096 0.0110 0.0133 0.0057 -0.0237 -0.0237 -0.0190 -0.0081 -0.0087 -0.0081 -0.0162 -0.0128 -0.0118 -0.0201 -0.0036 -0.0234 -0.0056 Sands Slickrock Upland 0.0184 0.0184 0.0023 -0.0556 -0.0556 -0.0174 -0.0253 -0.0049 -0.0223 -0.0318 -0.0439 -0.0491 -0.0108 -0.0578 -0.0053 -0.0214 -0.0435 -0.0227 -0.0135 -0.0014 Rocky Slopes 0.0135 0.0135 0.0214 Arid -0.0644 -0.0644 -0.0432 -0.0419 -0.0650 -0.0714 -0.0708 -0.0158 -0.0402 -0.0256 -0.0277 -0.0392 -0.0197 -0.0124 -0.0222 -0.0023 -0.0014 Lowlands 0.0108 0.0108 0.0232 0.0144 0.0062 0.0077 Flats -0.0087 -0.0087 -0.0133 -0.0206 -0.0183 -0.0105 -0.0169 -0.0225 -0.0110 -0.0004 -0.0075 -0.0020 -0.0040 -0.0036 Lowland

uegrass Semi-natural Shrubland uegrass Semi-natural Shrubland

5. Continued. PPENDIX A XXx. Alternative Classification HierarchyClassification Alternative XXx. Association NVC Quaking Aspen / Gambel Oak / Mountain Snowberry Forest Snowberry / Mountain Oak Aspen / Gambel Quaking Vegetation Semi-natural Wheatgrass Herbaceous Intermediate Bl / Kentucky Snowberry Mountain Shrubland Snowberry / Mountain Oak Gambel Forest Snowberry / Mountain Fir White Forest Snowberry / Mountain Fir Aspen - White Quaking Shrubland Serviceberry Utah Plateau / High B2. Streambank / Herbaceous Shrubland / Woodland / Forest - Plateau / High Streambank B2a. Vegetation Sedge Herbaceous Nebraska BalticHerbaceous RushVegetation Shrubland Birch Wooded / Water Cottonwood Fremont Sedge Shrubland / Beaked Willow Strapleaf WoodlandUnderstory / Disturbed Box-elder / Herb / Shrub / Woodland Forest - Streambank B2b. Shrubland Seepwillow Emory Shrubland Graminoids / Mesic Willow Coyote (Field Horsetail, Variegated Scouringrush) Herbaceous Vegetation Ash Woodland - Velvet Cottonwood Fremont Woodland / Emory Seepwillow Cottonwood Fremont Forest Willow / Coyote Cottonwood Fremont

84 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 us d / Herbaceous / Woodland / Herbaceous/ Woodland rubland / Herbaceousrubland dland / Shrubland / Herbaceous / Shrubland dland 2004), and their placement in alternative rest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland rest Crevice Canyon - Forest based on coalition group affinities - Forest / Woodland / Shrubland / Herbaceo / Shrubland / Woodland - Forest / Crevice Canyon - Forest / Woodland / Herbaceous / Woodland - Canyon Forest / Crevice / Herbaceous / Woodland - Canyon Forest / Crevice / Crevice Canyon - Forest / Woodland / Herbaceous / Woodland - Canyon Forest / Crevice u / Crevice Canyon - Forest / Woodland / Herbaceous / Woodland - Canyon Forest u / Crevice eau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland eau - Forest Mesa Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top -Mesa 2b. Streambank - Forest / Woo - Forest 2b. Streambank

B1b. High Plateau - Forest / Woodland / Sh / Woodland - Forest Plateau High B1b. ass Semi-natural Forest Forest ass Semi-natural Plateau High B1b. wberry Forest wberry / Herbaceous / Shrubland / Woodland - Forest Plateau High B1b. st B1a. High Plateau / Oak Forest Forest Oak / Woodlan - Canyon Forest / Crevice High B1a. Plateau / Mountain Snowberry / Tall Forbs Forest Forbs / Tall Snowberry / Mountain

Cottonwood / Coyote Willow Forest Forest Willow / Coyote Cottonwood B Forest Manzanita / Greenleaf Fir White Forest Oak / Gambel Fir White Forest Snowberry / Mountain Fir White Forest Oak / Gambel Douglas-fir Douglas-fir Snowberry / Mountain Fore A1a. Plat High B1b. High B1a. Plateau High B1a. Plateau Sno / Mountain Fir - Aspen White Quaking White Fir / Bigtooth Maple Forest / Bigtooth Fir White Maple Forest / Bigtooth Douglas-fir High B1a. Plateau High B1a. Platea Bluegr / Kentucky Fir - Aspen White Quaking liance Forest Snowberry / Mountain Oak / Gambel Aspen Quaking - Fo Plateau High B1b. 6. Vegetation units of the U.S. National Vegetation Classification at Zion National Park, Utah, as recognized by Cogan et al. ( Classification at Zion National Park, National Vegetation units of the U.S. 6. Vegetation PPENDIX A Alliance Association Alternative class X. CLASS X.X. Subclass GroupX.X.X. Formation Alternative X.X.X.X.x. Formation Alliance Association I. FOREST forest I.A. evergreen forest evergreen needle-leaved I.A.8. forest evergreen needle-leaved or subpolar temperate Conical-crowned I.A.8.N.c. Alliance Forest Fir White Alliance Forest Douglas-fir forest I.B. deciduous forest cold-deciduous I.B.2. forest cold-deciduous boreal or Montane I.B.2.N.b. Alliance Forest Maple Montane Bigtooth Alliance Forest Aspen Quaking forest cold-deciduous flooded Temporarily I.B.2.N.d. Al Forest Flooded Aspen Temporarily Quaking Alliance Forest Flooded Temporarily Cottonwood Fremont forest - deciduous evergreen mixed I.C. forest - cold-deciduous evergreen needle-leaved mixed I.C.3. forest - cold-deciduous evergreen needle-leaved Mixed I.C.3.N.a. Fremont Maple / Gambel Bigtooth Alliance Forest Aspen - Quaking Fir White Aspen Quaking classes based on coalition group affinities (see Fig. 7 in paper). classes based on coalition group affinities (see Fig. 2015] PLANT COALITION GROUPS OF ZION NP 85 Woodland / Shrubland / / Shrubland Woodland / / Shrubland Woodland st / Woodland / Herbaceous / Woodland st rest / Woodland / Herbaceous / Woodland rest y Slopes - y Slopes - odland / Shrubland / Sparse / Sparse / Shrubland odland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland / Sparse / Shrubland Woodland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland land / Shrubland / Sparse / Sparse / Shrubland land land / Shrubland / Sparse / Sparse / Shrubland land oodland / Shrubland / Shrubland oodland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland based on coalition group affinities - Woodland / Shrubland - / Shrubland Woodland - / Shrubland Woodland es - Woodland / Shrubland - / Shrubland es Woodland ateau / Crevice Canyon - Forest / Woodland / Herbaceous / Woodland - Canyon Forest / Crevice ateau Top / Rocky Slopes - Woodland / Shrubland / Sparse / Sparse - / Shrubland Slopes Woodland Top / Rocky Plateau / Crevice Canyon - Canyon Fo / Crevice Plateau Mesa Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top -Mesa 1a. Mesa Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top - 1a. Mesa 1b. Mesa Top / Rocky Slopes - Wo Slopes Top / Rocky 1b. Mesa

A1c. Upland Sands / Slickrock / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse - Slopes Top / Rocky A1b. Mesa unavailable) data (plot unclassified A2d. Rocky Slopes - Slopes W A2d. Rocky - Slopes Top / Rocky A1b. Mesa - Slopes Top / Rocky A1b. Mesa - Slopes Top / Rocky A1b. Mesa land land / Shrubland / Woodland Forest Top - A1a. Mesa nd - Wood Lowlands A2b. Arid - Wood Lowlands A2b. Arid Woodland Slopes A2d. Rocky d Woodland d Woodland unavailable) data (plot unclassified Woodland Woodland Slopes A2d. Rocky Oak WoodlandOak Slop A2d. Rocky X.X.X.X.x. Formation. Woodland Woodland - Lowlands A2b. Arid  r - r Oak Gambel Woodland - Canyon Fore / Crevice High B1a. Plateau

Mountain-mahogany / Littleleaf - Juniper Utah Two-needle Pinyon Woodland Woodland Cliff-rose / Stansbury - Juniper Utah Pinyon Two-needle Woodland Sagebrush / Big species - Juniper Pinyon Two-needle / Mountain-mahogany species Woodland - Juniper Pinyon Two-needle Woodland Oak / Gambel species - Juniper Pinyon Two-needle Wood Mountain-mahogany / Curl-leaf Pinyon Two-needle A Woodland Sagebrush / Black Pine Ponderosa Woodland / Smooth Semi-natural Brome Pine Ponderosa Woodland Bracken / Northern Pine Ponderosa Woodland Oak / Gambel Pine Ponderosa High B1a. Sagebrush / Black - Juniper Utah Pinyon Singleleaf A Woodla Sagebrush / Big - Juniper Utah Pinyon Singleleaf High B1a. Pl Oak / Mountain-mahogany- - Juniper Gambel Utah Pinyon Singleleaf Woodland Woodland / Blackbrush - Juniper Utah Pinyon Singleleaf A1a. Galleta / James' / Snakeweed - Juniper Utah Pinyon Singleleaf Woodland / Needle-and-Threa - Juniper Utah Pinyon Singleleaf Live / Turbinella - Juniper Utah Pinyon Singleleaf Understory / Sparse - Juniper Utah Pinyon Singleleaf Two-needle Pinyon - Utah Juniper / Greenleaf Manzanita Woodland Manzanita / Greenleaf - Juniper Utah Pinyon Two-needle Manzanita Woodland / Greenleaf Pine Ponderosa A1b. Mesa Utah Buffaloberry, / (Roundleaf - Juniper Utah Pinyon Singleleaf Serviceberry) Woodland X.X.X. Formation Group X.X.X. Formation  X.X. Subclass  6. Continued. X. Class PPENDIX A Alliance Association Alternative class Alternative Alliance Association II. WOODLAND woodland II.A. evergreen woodland evergreen needle-leaved II.A.4. woodland evergreen needle-leaved or subpolar temperate Rounded-crowned II.A.4.N.a. Alliance species) Woodland - (Juniper Pinyon Two-needle Alliance Woodland Pine Ponderosa Alliance Woodland Juniper Utah Singleleaf Pinyon - (Utah Juniper) WoodlandAlliance / Big Sagebrush Juniper Utah Alliance Woodland Mountain Juniper Rocky woodland II.B. deciduous woodland cold-deciduous II.B.2. woodland Cold-deciduous II.B.2.N.a. Semi-natural WoodlandRussian-olive Alliance Junipe Mountain Rocky Semi-natural Woodland Russian-olive unavailable) (plot data unclassified 86 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 Woodland / Shrubland / / Shrubland Woodland Woodland / Shrubland / / Shrubland Woodland / / Shrubland Woodland Woodland / Shrubland / / Shrubland Woodland / Shrubland / Herbaceous / Shrubland y Slopes - y Slopes - y Slopes - y Slopes - Shrubland / Sparse Shrubland oodland / Shrubland / Sparse / Sparse / Shrubland oodland Forest / Woodland / Shrubland / / Shrubland / Woodland Forest / Shrubland / Sparse / Sparse / Shrubland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland nd / Shrubland / Sparse / Sparse nd / Shrubland nd / Shrubland nd / Shrubland Lowland Flats - Woodland / Shrubland / - / Shrubland Lowland Flats Woodland based on coalition group affinities bank - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -bank Forest Slopes -Slopes Woodla Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top - Plateau / Crevice Canyon - Forest / Woodland / Herbaceous / Woodland - Canyon Forest / Crevice Plateau Lowlands - Woodland - Woodland Lowlands ambank - Forest / Woodland - Forest ambank 2d. Rocky Slopes - Slopes 2d. Rocky Woodland / Shrubland Herbaceous / Sparse A2a. Arid Lowlands / Lowlands A2a. Arid Herbaceous / Sparse / Rock / Slickrock Sands A1c. Upland

B2a. Streambank / High Plateau - Plateau / High Streambank B2a. - Slopes Top / Rocky A1b. Mesa / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse Herbaceous A1c. Upland Sands / Slickrock / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse - Woodla Lowlands A2b. Arid / Shrubland / Woodland Forest Top - A1a. Mesa land A2d. Rocky Rocky land A2d. and unclassified (plot data unavailable) data (plot and unclassified X.X.X.X.x. Formation. rass Semi-natural rass Shrubland - Slopes Sands / Rocky A2c. Upland  (Utah Serviceberry) Colluvial Shrub Colluvial Serviceberry) (Utah brush Colorado Plateau Shrubland Shrubland Plateau Colorado brush A2b. Arid

untain Big Sagebrush / Needle-and-Thread Sagebrush Big Shrubland untain - Slopes W Top / Rocky A1b. Mesa

Woodland Understory / Disturbed Box-elder Woodland Seepwillow / Emory Cottonwood Fremont Stre B2b. - Shrubland (Utah Oak - Gambel Serviceberry) Manzanita Greenleaf Shrubland Manzanita Greenleaf Shrubland Grama / Blue Sagebrush Big Salt-cedar species Temporarily FloodedShrubl Box-elder / Tasselflower Brickelbush Brickelbush Woodland / Tasselflower Box-elder Ash Woodland - Velvet Cottonwood Fremont High B1a. Stream B2b. - Mountain Shrubland Big Manzanita Sagebrush Greenleaf Shrubland Manzanita Mexican Semi-natural / Cheatgrass Rabbitbrush) - Sagebrush (Rubber Big Shrubland Rye) Wild - (Streamside Wheatgrass / Western Sagebrush Big Basin Shrubland A1a. Mesa

X.X.X. Formation Group X.X.X. Formation  ance Shrubland - Manzanita Cliff-rose Greenleaf Stansbury X.X. Subclass nd Alliance Single-leaf Ash Woodland A  ed Woodland Alliance 6. Continued. X. Class PPENDIX A Alliance Association Alternative class Alternative Alliance Association woodland cold-deciduous flooded Temporarily II.B.2.N.b. Box-elder Temporarily Flood Alliance Woodland Flooded Temporarily Cottonwood Fremont Single-leaf Ash Temporarily Flooded Woodla III. SHRUBLAND shrubland evergreen III.A. shrubland evergreen broad-leaved III.A.2. shrubland evergreen broad-leaved temperate Sclerophyllous III.A.2.N.c. Greenleaf Manzanita Shrubland Alliance Shrubland Alliance Manzanita Mexican Alliance Shrubland Oak Live Turbinella shrubland evergreen broad-leaved temperate flooded Seasonally III.A.2.N.h. Arrow-weed Seasonally Flooded Shrubland Alliance shrubland evergreen microphyllous III.A.4. shrubland evergreen microphyllous Lowland III.A.4.N.a. Big Sagebrush Shrubland Alliance Arrow-weed Seasonally Flooded Shrubland - Oak Live Turbinella Basin Big Sagebrush Shrubland Alliance Alliance Sagebrush Big Shrubland Mountain Alli Cliff-rose) Shrubland Mexican Cliff-rose, (Stansbury Alliance Shrubland Rabbitbrush Rubber Alliance Shrubland Sand Sagebrush shrubland microphyllous flooded Temporarily III.A.4.N.c. Alliance Flooded Shrubland Temporarily Semi-natural species Salt-cedar Mo / Cheatg Rabbitbrush Rubber Sand Sage 2015] PLANT COALITION GROUPS OF ZION NP 87

nd / Shrubland / nd / Shrubland Woodland / Shrubland / / Shrubland Woodland nd / Shrubland / nd / Shrubland nd / Shrubland / Sparse y Slopes - Woodla y Slopes - Forest / Woodland / Shrubland / / Shrubland / Woodland Forest / / Shrubland / Woodland Forest / Shrubland / Sparse / Sparse / Shrubland / Woodland / Shrubland / Herbaceous / Shrubland / Woodland wland Flats - Woodland / Shrubland / / -Shrubland Flats Woodland wland Lowland Flats -Lowland Flats Woodla Lowland Flats - Woodland / Shrubland / - / Shrubland Lowland Flats Woodland based on coalition group affinities ds - Woodland / Shrubland / Sparse / Sparse / Shrubland -ds Woodland p / Rocky Slopes -p / Rocky Slopes Woodla bank - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -bank Forest ateau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -ateau Forest Top / Rocky Slopes - Woodland / Shrubland / Sparse / Sparse - / Shrubland Slopes Woodland Top / Rocky Lowlands - Woodland Lowlands ands / Rocky Slopes / Upland Sands (outlier) Slopes / Rocky ands Herbaceous / Sparse B2a. Streambank / High Plateau - Plateau / High Streambank B2a. Herbaceous - Plateau / High Streambank B2a. Herbaceous

Herbaceous / Sparse A2a. Arid Lowlands / Lowlands A2a. Arid / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse / Rock / Slickrock Sands A1c. Upland A2a. Arid Lowlands / Lowlands A2a. Arid Herbaceous / Sparse Herbaceous / Sparse A2a. Arid Lowlands / Lo Lowlands A2a. Arid ural Shrubland Shrubland ural - Forest Plateau High B1b. dge) Shrubland A1b. Mesa To X.X.X.X.x. Formation.  Basalt ShrublandBasalt

in Snowberry / Kentucky Bluegrass / Kentucky Semi-nat Snowberry in Serviceberry Shrubland Serviceberry / Herbaceous / Shrubland / Woodland - Forest Plateau High B1b. Shrubland Blackbrush Shrubland Saltbush Fourwing Shrubland Serviceberry / Utah Oak Gambel Shrubland Sagebrush / Big Oak Gambel Lowlan A2b. Arid Shrubland / Muttongrass Oak Gambel Shrubland Snowberry / Mountain Oak Gambel A1b. Mesa Shrubland Graminoids / Mesic Willow Coyote Pl High B1b. Stream B2b. Nevada Joint-fir Shrubland / Galleta James' Blackbrush Shrubland - Sagebrush Big Saltbush Fourwing A2b. Arid -Gambel Oak Mountain-mahogany(Geyer's Se / Shrubland / Barren Willow Coyote A2. Lowl X.X.X. Formation Group X.X.X. Formation  X.X. Subclass  Temporarily Shrubland Flooded 6. Continued. X. Class PPENDIX A Alliance Association Alternative class Alternative Alliance Association III.A.5. extremely xeromorphic subdesert shrubland shrubland subdesert xeromorphic extremely III.A.5. shrubland subdesert xeromorphic extremely evergreen microphyllous and Broad-leaved III.A.5.N.a. Nevada Joint-fir Shrubland Alliance Mormon-tea ShrublandAlliance shrubland subdesert xeromorphic extremely deciduous Facultatively III.A.5.N.b. Alliance Shrubland Blackbrush Alliance Shrubland Saltbush Fourwing shrubland deciduous III.B. shrubland cold-deciduous III.B.2. shrubland cold-deciduous Temperate III.B.2.N.a. Shrubland Alliance Gray Horsebrush Snowberry Mountain - Mormon-tea Shrubland Alliance Shrubland Serviceberry Utah Alliance Shrubland Oak Gambel A2c. Upland Sands / Rocky Slopes - Shrubland / Sparse Mounta shrubland cold-deciduous flooded Temporarily III.B.2.N.d. Water BirchTemporarily Flooded Shrubland Alliance Utah Strapleaf Willow Temporarily Flooded Shrubland Alliance (Coyote Willow, Sandbar Willow) Alliance Fremont Cottonwood / Water Birch WoodedShrubland Strapleaf Willow / Beaked Sedge Shrubland 88 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 Woodland / Shrubland / / Shrubland Woodland / / Shrubland Woodland Woodland / Shrubland / / Shrubland Woodland / Woodland / Herbaceous / Woodland y Slopes - y Slopes - y Slopes - Forest / Woodland / Shrubland / / Shrubland / Woodland Forest / / Shrubland / Woodland Forest lickrock / Rocky Slopes (outlier) dland / Shrubland / Sparse / Sparse / Shrubland dland Lowland Flats - Woodland / Shrubland / - / Shrubland Lowland Flats Woodland / - / Shrubland Lowland Flats Woodland based on coalition group affinities u / Crevice Canyon - Forest lateau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland - Forest lateau Top / Rocky Slopes - Woodland / Shrubland / Sparse / Sparse - / Shrubland Slopes Woodland Top / Rocky Lowlands - Woo Lowlands Upland Sands / Rocky Slopes - Shrubland / Sparse - Slopes Shrubland / Rocky Sands Upland Herbaceous / Sparse Herbaceous / Sparse A1c. Upland Sands / Slickrock / Rock / Slickrock Sands A1c. Upland / Herbaceous / Shrubland / Woodland - Forest Plateau High B1b. / Lowlands A2a. Arid Herbaceous / Sparse / Lowlands A2a. Arid Herbaceous / Sparse / Rock / Slickrock Sands A1c. Upland - Plateau / High Streambank B2a. Herbaceous / Herbaceous / Shrubland / Woodland - Forest Plateau High B1b. - Plateau / High Streambank B2a. Herbaceous

A1c. Upland Sands / Slickrock / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse bland A2b. Arid Arid bland A2b. Seasonally Flooded Herbaceous Flooded Herbaceous Seasonally ies) / James' Galleta Dwarf-shru / James'ies) Galleta X.X.X.X.x. Formation. hread Herbaceous Vegetationhread Herbaceous  Basin Herbaceous Basin Vegetation Herbaceous Vegetation ate Wheatgrass Semi-natural Herbaceous Vegetation Herbaceous Semi-natural Wheatgrass ate HighP B1b. Sedge Herbaceous Vegetation

(Sandhill Muhly, Mountain Muhly) - Hairy Goldenaster Herbaceous Vegetation Herbaceous Semi-natural Wheatgrass) Brome Smooth - (Western Vegetation Vegetation Herbaceous Galleta James' Semi-natural Bluegrass Kentucky Alliance SedgeBeaked Herbaceous Vegetation B1a. High Platea Sagebrush / Needle-and-Thread Dwarf-shrubland Black Dwarf-shrubland / Muttongrass Sagebrush Black A1b. Mesa Black Sagebrush / Bottlebrush Dwarf-shrubland / Bottlebrush Sagebrush Black A2c. X.X.X. Formation Group X.X.X. Formation  X.X. Subclass  Alliance Baltic Rush Alliance Baltic Seasonally Flooded Herbaceous Herbaceous Flooded Seasonally Seasonally Flooded Herbaceous Flooded Herbaceous Seasonally ed Herbaceous Allianceed Herbaceous Nebraska e Blue Grama - Needle-and-T - Grama e Blue liance Sand Dropseed Great Dropseed liance Sand 6. Continued. X. Class PPENDIX A Alliance Association Alternative class Alternative Alliance Association Alliance Alliance IV.B. deciduous dwarf-shrubland dwarf-shrubland deciduous IV.B. dwarf-shrubland cold-deciduous IV.B.2. dwarf-shrubland cold-deciduous Caespitose IV.B.2.N.a. Alliance Dwarf-shrubland Snakeweed VEGETATION V. HERBACEOUS grassland perennial V.A. grassland subpolar or temperate V.A.5. grassland or subpolar sod temperate Medium-tall V.A.5.N.c. Alliance Semi-natural Herbaceous Wheatgrass Intermediate grassland or subpolar bunch temperate Medium-tall V.A.5.N.d. Alliance MuhlyMountain Herbaceous Intermedi Needle-and-Thread Bunch HerbaceousAlliance spec - Snakeweed (Prickly-pear Alliance Semi-natural Smooth Brome Herbaceous Sand Dropseed Herbaceous Al grassland subpolar or temperate sod Short V.A.5.N.e. Alliance Herbaceous Galleta James' Needle-and-Thread Great Basin Herbaceous Vegetation Blue Grama Herbaceous Allianc grassland or subpolar temperate flooded Seasonally V.A.5.N.k. HerbaceousSeasonallyFlooded Baltic Rush Semi-natural Bluegrass Kentucky A1. Mesa Top / UplandSands / S Alliance Sedge) Sedge, Beaked (Swollen-beak Nebraska Sedge Seasonally Flood IV. DWARF-SHRUBLAND IV. DWARF-SHRUBLAND dwarf-shrubland evergreen IV.A. dwarf-shrubland subdesert evergreen xeromorphic extremely IV.A.2. dwarf-shrubland subdesert evergreen xeromorphic Extremely IV.A.2.N.a. Alliance Dwarf-shrubland Sagebrush Black 2015] PLANT COALITION GROUPS OF ZION NP 89 s Woodland / Shrubland / / Shrubland Woodland Woodland / Shrubland / / Shrubland Woodland y Slopes - y Slopes - Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland nd / Shrubland / Sparse / Sparse nd / Shrubland rest / Woodland / Shrubland / Herbaceous / Woodland rest Lowland Flats - Woodland / Shrubland / - / Shrubland Lowland Flats Woodland based on coalition group affinities Forest / Woodland / Shrubland / Herbaceou / Shrubland / Woodland Forest ands / Rocky Slopes - Shrubland / Sparse - Slopes Shrubland / Rocky ands p / Rocky Slopes -

A2. Lowlands / Rocky Slopes / Upland Sands (outlier) Slopes / Rocky A2. Lowlands A2a. Arid Lowlands / Lowlands A2a. Arid Herbaceous / Sparse A1c. Upland Sands / Slickrock / Rock / Slickrock Sands A1c. Upland Herbaceous / Sparse

A2b. Arid Lowlands - Woodla Lowlands A2b. Arid Herbaceous / Sparse A1c. Upland Sands / Slickrock / Rock / Slickrock Sands A1c. Upland getation A1b. Mesa To Herbaceous Vegetation B2b. Streambank - Fo X.X.X.X.x. Formation.  ck Pavement Sparse Ve ck Pavement s Semi-natural Alliance s unavailable) data (plot unclassified ahogany Slickrock Vegetation Sparse en Sparse Vegetation en ory Seepwillow Shrubland Shrubland Seepwillow ory - Streambank B2b. bber Rabbitbrush Sand Deposit Sparse Vegetation Vegetation Sparse Deposit Sand Rabbitbrush bber S A2c. Upland

Green Rabbitbrush / Kentucky Bluegrass Semi-natural Shrub Shrub Semi-natural Bluegrass / Kentucky Rabbitbrush Green Herbaceous Vegetation (Field Horsetail, Variegated Scouringrush) Vegetation Sparse Badlands Buckwheat Wild Crispleaf Nevada Joint-fir / Lich X.X.X. Formation Group X.X.X. Formation  X.X. Subclass  getated Alliance Littleleaf Mountain-m ted Alliance Mountain-mahogany Ro cular vegetationcular with sparse vas with sparse 6. Continued. X. Class PPENDIX A Alliance Association Alternative class Alternative Alliance Association V.A.7. temperate or subpolar grassland with a sparse needle-leaved or microphyllous evergreen shrub layer layer shrub evergreen microphyllous or needle-leaved a sparse with grassland subpolar or temperate V.A.7. layer shrub evergreen microphyllous or needle-leaved a sparse with subpolar grassland or temperate Medium-tall V.A.7.N.e. Herbaceous Shrub Rabbitbrush Green Alliance vegetation forb perennial V.B. forb vegetation perennial temperate V.B.2. forb vegetation perennial temperate flooded Semipermanently V.B.2.N.e. Flooded Semipermanently Scouringrush) Variegated Horsetail, (Field Alliance Herbaceous V.D. annual grassland grassland V.D. annual grassland annual V.D.2. temperate grassland annual temperate Short V.D.2.N.d. Alliance Semi-naturalCheatgrass Herbaceous VEGETATION VII. SPARSE vegetation vascular sparse VII.A. vegetation vascular sparse with cliffs VII.A.1. VII.A.1.N.a. Cliffs Alliance Vegetated Sparsely Bedrock Wooded vegetation vascular sparse with pavement VII.A.2. Herbaceou Cheatgrass vegetation vascular sparse with Pavement VII.A.2.N.a. Littleleaf Mountain-mahogany Ve Sparsely slopes VII.C. slopes dry VII.C.3. Vegetation Sparse Slickrock Pine Ponderosa slopes Dry VII.C.3.N.b. Mountain-mahogany Vegeta Sparsely Alliance Vegetated Sparsely Desert Painted UNDETERMINED PLACEMENT HIERARCHY [provisional] Shrubland Seepwillow Emory [provisional] Sparse Vegetation Sand Deposit Rabbitbrush Rubber Ru Em

90 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 us according to the revised classification Woodland / HerbaceousWoodland / HerbaceousWoodland / Woodland / Herbaceous/ Woodland / Herbaceous/ Woodland / Herbaceous/ Woodland C classes were obtained or inferred from NatureServe - / Herbaceous / Woodland Forest oodland / Shrubland / / HerbaceousShrubland oodland / Woodland / Shrubland / Herbaceous / Shrubland / Woodland / Woodland / Shrubland / Herbaceous / Shrubland / Woodland ice Canyon - / Forest ice Canyon - / Forest ed on coalition group affinities Crevice Canyon - Forest - Forest / Woodland / Shrubland / Herbaceo / Shrubland / Woodland - Forest / Crevice Canyon - Forest / Crevice Canyon - Forest High Plateau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -High Forest Plateau Mesa Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top -Mesa B1a. High / Crev Plateau B1a. High / Crev Plateau B1a. High / Crevice Plateau Canyon

menziesii Mesic Southern Rocky Mountain Forest Group Group Forest Mountain Rocky Southern Mesic menziesii ass Semi-natural Forest Forest ass Semi-natural High B1b. Plateau Forbs Forest Forbs - High Forest B1b. Plateau wberry Forest wberry - High Forest B1b. Plateau esii Southern Rocky Mountain Dry Forest Group Forest Dry Mountain Rocky Southern esii st B1a. High Plateau / st / W - High Forest B1b. Plateau cky Mountain Montane Forest & Woodland Group Woodland & Forest Montane cky Mountain & High Montane Conifer Forest Forest Conifer Montane High & cky Mountain Forest & Woodland Group Group Woodland & Forest Mountain cky 7. Associations of the U.S. National Vegetation Classification (NVC) at Zion National Park, Utah (Cogan et al. 2004), arranged Classification (NVC) at Zion National Park, National Vegetation 7. Associations of the U.S. ass l PPENDIX A X - C Woodland Open to 1 - Forest Forest Boreal & - Temperate 1.B Forest Temperate - Cool 1.B.2 Forest Temperate Cool Mountain Rocky - 1.B.2.Nb Forest Foothill & Montane Dry Lower Mountain Rocky Central Ro Middle menziesii Pseudotsuga Douglas-fir Snowberry / Mountain Fore Subalpine Rocky Mountain Ro tremuloides Populus / Tall Snowberry Aspen / Mountain Quaking Forest Snowberry / Mountain Oak Aspen / Gambel Quaking Forest Montane Lower Mountain Rocky Southern Pseudotsuga - pungens Picea - concolor Abies Bigtooth Maple Forest / Gambel Oak Douglas-firMaple / Bigtooth Forest Bluegr / Kentucky Fir Aspen - White Quaking Sno / Mountain Fir - Aspen White Quaking MapleWhite Fir / Bigtooth Forest B1b. Fore Snowberry / Mountain Fir White menzi Pseudotsuga - concolor Abies Douglas-fir Forest / Gambel Oak Forest Fir / Gambel Oak White Forest Manzanita / Greenleaf Fir White B1a. High Plateau A1a. B1a. High Plateau X.X - Subclass X.X.X - Formation X.X.X.Xx - Division Macrogroup Group Association class Alternative bas heirarchy (FGDC 2008), and their placement in alternative classes based on coalition group affinities (see Fig. 7 in paper). NV heirarchy (FGDC 2008), and their placement in alternative classes based on coalition group affinities (see Fig. (2013) or USNVC (2013). 2015] PLANT COALITION GROUPS OF ZION NP 91 d / Herbaceous / Sparse d / Herbaceous d / Shrubland / Herbaceous / Sparse / Sparse / Sparse land / Herbaceousland Woodland / HerbaceousWoodland d land / Shrubland / Sparse / Sparse / Shrubland land odland / Shrubland / Sparse / Sparse / Shrubland odland Group. on - / Wood Forest Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland / Sparse / Shrubland Woodland  land / Shrubland / Sparse / Sparse / Shrubland land land / Shrubland / Sparse / Sparse / Shrubland land ice Canyon - / Forest Slopes - Woodland / Shrubland / Shrubland - Slopes Woodland ed on coalition group affinities - Woodland / Shrubland - / Shrubland Woodland - Woodland / Shrubland - / Shrubland Woodland es - Woodland / Shrubland - / Shrubland es Woodland Macrogroup  ateau / Crevice Canyon - Forest / Woodland / Herbaceous / Woodland - Canyon Forest / Crevice ateau Top / Rocky Slopes - Slopes Wood Top / Rocky y Slopes - / Shrublan Slopes y Woodland Lowlands - Woodland / Shrubland / Sparse / Sparse / Shrubland - Woodland Lowlands Mesa Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top -Mesa Upland Sands / Slickrock / Rocky Slopes - / Rocky / Shrublan Slopes Woodland / Slickrock Sands Upland 1b. Mesa Top / Rocky Slopes - Slopes Wo Top / Rocky 1b. Mesa 1a. Mesa Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top -1a. Mesa X.X.X.Xx - Division  odland odland / Rocky - Woodlan Slopes / Slickrock Sands A1c. Upland - Slopes Top / Rocky A1b. Mesa - Slopes Top / Rocky A1b. Mesa k Woodland Woodland k Top / Rocky A1b. Mesa - Slopes Top / Rocky A1b. Mesa Serviceberry) Woodland A2d. Rock X.X.X - Formation X.X.X - Formation land land / Shrubland / Woodland Forest Top - A1a. Mesa  nd A2b. Arid Lowlands - Wood - Lowlands Arid nd A2b. A2b. Arid Lowlands - Wood - Lowlands Arid A2b. Woodland Slopes A2d. Rocky Woodland Woodland Slopes A2d. Rocky Oak WoodlandOak Slop A2d. Rocky oodland oodland B1a. High / Crevice Cany Plateau X.X Subgrass  ah Juniper - Western Juniper Woodland Woodland Juniper - Western ah Juniper mi-natural Woodlandmi-natural B1a. High / Crev Plateau Rocky Mountain Forest & Woodland Group Group Woodland & Forest Mountain Rocky 7. Continued. X - Class PPENDIX A Association class Alternative bas 1.B.2.Nc - Western North American Cool Temperate Woodland & Scrub Scrub & Woodland Temperate Cool American North Western - 1.B.2.Nc - Pinyon Singleleaf Ut Intermountain Group Savanna & Woodland osteosperma Juniperus Rocky Mountain Juniper - Gambel Oak W Woodland / Big Sagebrush Juniper Utah Group Woodland osteosperma - Juniperus edulis Pinus Woodland Sagebrush / Big species - Juniper Pinyon Two-needle Woodland Oak / Gambel species - Juniper Pinyon Two-needle / Mountain-mahogany species Woodland - Juniper Pinyon Two-needle Woodland Manzanita / Greenleaf - Juniper Utah Pinyon Two-needle Wo Mountain-mahogany / Littleleaf - Juniper Utah Pinyon Two-needle Woodland Cliff-rose / Stansbury - Juniper Utah Pinyon Two-needle Wood Mountain-mahogany / Curl-leaf Pinyon Two-needle A1c. Group Woodland osteosperma - Juniperus monophylla Pinus Utah Buffaloberry, / (Roundleaf - Juniper Utah Pinyon Singleleaf A Woodla Sagebrush / Big - Juniper Utah Pinyon Singleleaf Sagebrush / Black - Juniper Utah Pinyon Singleleaf A2b. Arid Woodland / Blackbrush - Juniper Utah Pinyon Singleleaf / Mountain-mahogany- Oa - Juniper Gambel Utah Pinyon Singleleaf Understory / Sparse - Juniper Utah Pinyon Singleleaf Live / Turbinella - Juniper Utah Pinyon Singleleaf Pinus ponderosa Southern Southern ponderosa Pinus Woodland Sagebrush / Black Pine Ponderosa Woodland Oak / Gambel Pine Ponderosa Manzanita Woodland / Greenleaf Pine Ponderosa Woodland Bracken / Northern Pine Ponderosa / SmoothPonderosa Pine Brome Se A A1b. Mesa A1a. High B1a. Pl 92 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 Herbaceous / Sparse rubland / Herbaceous / Sparserubland and / Shrubland / Herbaceous / Sparse land / Herbaceousland oodland / Shrubland / Herbaceous / Shrubland oodland / Woodland / Shrubland / Herbaceous / Shrubland / Woodland nd / Shrubland / Sparse lopes - Woodland / Shrubland / lopes / Shrubland - Woodland / Shrubland / Herbaceous / Shrubland / Shrubland / Herbaceous / Shrubland cky - Slopes / Sh Woodland Group. on - / Wood Forest  ed on coalition group affinities - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland - Forest / High Plateau - Forest / W - Forest Plateau / High Macrogroup nk - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -nk Forest p / Rocky Slopes -p / Rocky Slopes Woodla  bank - Forest / Woodland - Forest bank ateau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -ateau Forest y Slopes - Woodland / Shrubland - / Shrubland Slopes y Woodland ambank - Forest / Woodland -ambank Forest land Sands / Slickrock / Rocky S / Slickrock Sands land Upland Sands / Slickrock Ro B2a. Streambank / High Plateau - Forest Plateau / High Streambank B2a.

X.X.X.Xx - Division  X.X.X - Formation X.X.X - Formation  virginiana Southern Rocky Mountain Montane Shrubland Group Group Shrubland Montane Mountain Rocky Southern virginiana dge) Shrubland A1b. Mesa To nsis - Rhus trilobata Foothill Shrubland Group Group Shrubland Foothill trilobata - Rhus nsis wrightii - Juglans major Warm Desert Riparian Scrub Group Group Scrub Riparian Desert Warm major - Juglans wrightii rubra - Alnus rhombifolia Riparian Woodland Group Group Woodland Riparian rhombifolia - Alnus rubra X.X Subgrass - Muhlenbergia montana Grassland Group Group Grassland montana Muhlenbergia - oodland oodland B1a. High / Crevice Cany Plateau  7. Continued. X - Class PPENDIX A Association class Alternative bas 1.B.3 - Temperate Flooded & Swamp Forest Forest & Swamp Flooded - Temperate 1.B.3 Forest Swamp Flooded & Basin & Great Mountain Rocky -1.B.3.Nc Forest Rocky Mountain & Great Basin Riparian Alnus - wislizeni ssp. deltoides Populus Woodland Understory / Disturbed Box-elder Box-elder / Tasselflower Brickelbush W Forest Willow / Coyote Cottonwood Fremont Ash Woodland Single-leaf Shrubland Birch Wooded / Water Cottonwood Fremont Forest Swamp & Flooded American North Southwestern -1.B.3.Nd Forest Riparian Southwest Warm Platanus - wislizeni ssp. deltoides Populus Ash Woodland - Velvet Cottonwood Fremont Woodland Seepwillow / Emory Cottonwood Fremont Streambank B2a. Streamba B2b. Stre B2b. A2d. Rock Stream B2b. 2 - Shrubland & Grassland Grassland & 2 - Shrubland & Shrubland Grassland Boreal & - Temperate 2.B & Shrubland Meadow Grassland, -2.B.2 Temperate & Grassland Shrubland 2.B.2.Na North American - Western Meadow Forb & Grass Mesic Montane High & Subalpine Mountain-Vancouverian Rocky thurberi Festuca - arizonica Festuca (Sandhill Muhly, Mountain Muhly) - Hairy Goldenaster Herbaceous Vegetation Shrubland Montane Mountain Southern Rocky utahe Amelanchier - montanus Cercocarpus Vegetation Mountain-mahogany Sparse Slickrock Littleleaf A1c. Upland Sands / Slickrock / Rocky Slopes - Woodl Shrubland Serviceberry Utah Prunus - spp. - Amelanchier gambelii Quercus -Gambel Oak Mountain-mahogany(Geyer's Se / Sagebrush Shrubland / Big Oak Gambel Shrubland Snowberry / Mountain Oak Gambel A1c. Up High B1b. Pl High B1b. Plateau A1c. 2015] PLANT COALITION GROUPS OF ZION NP 93 aceous / Sparse rbaceous / Sparse rubland / Herbaceous / Sparserubland aceous aceous odland / Shrubland / Herbaceousodland / Sparse land / Shrublandland / Herbaceous - Woodland / Shrubland / Herb / Shrubland - Woodland oodland / oodland Shrubland / Herbaceous / Woodland / Herbaceous / Woodland cky - Slopes / Sh Woodland Group. Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland  / Rocky Slopes - Woodland / Shrubland / He / Shrubland -/ Rocky Slopes Woodland ed on coalition group affinities gh Plateau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland - Forest Plateau gh High Plateau -Plateau / Wood Forest High - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland - Forest - Woodland / Shrubland - / Shrubland Woodland u / Crevice Canyon - Forest Macrogroup nk / High Plateau - / W Plateau Forest / High nk  Sands / Slickrock / Rocky Slopes / Slickrock Sands Sands / Slickrock Rocky - Slopes Wo Top / Rocky Slopes - Woodland / Shrubland / Sparse / Sparse / Shrubland - Slopes Woodland Top / Rocky Top - Forest / Woodland / Shrubland / Shrubland / Woodland Forest Top - Plateau - Forest / Woodland / Shrubland / Herb/ Shrubland -Plateau / Woodland Forest land Sands / Slickrock Ro High Plateau - Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland -High Forest Plateau A1b. Mesa Top / Rocky Slopes - Slopes Top / Rocky A1b. Mesa X.X.X.Xx - Division  tion / Herb / Shrubland / Woodland - High Forest B1b. Plateau X.X.X - Formation X.X.X - Formation  Shrubland Slopes A2d. Rocky ndula - Pascopyrum smithii Mixedgrass Prairie Group Group Prairie Mixedgrass smithii - Pascopyrum ndula er Shrubland, Wet Meadow & Marsh Marsh & Meadow Wet Shrubland, er h Shrubland h Shrubland / Slickrock Sands A1c. Upland X.X Subgrass  eggii - Arctostaphylos pungens Chaparral Group Group Chaparral pungens Arctostaphylos - eggii uegrass Semi-natural Shrubland uegrass Shrubland Semi-natural High B1b. Plateau Seasonally Flooded Herbaceous Alliance B1b. High e Shrubland B2a. Streamba ctostaphylos nevadensis - Ceanothus velutinus Montane Sclerophyll Scrub Group Group Scrub Sclerophyll Montane velutinus - Ceanothus nevadensis ctostaphylos Vegetation B2a. Streambank / 7. Continued. X - Class PPENDIX A Association class Alternative bas Gambel Oak / Muttongrass Shrubland / Muttongrass Oak Gambel Shrubland Serviceberry / Utah Oak Gambel - Shrubland (Utah Oak - Serviceberry) Gambel Manzanita Greenleaf Bl / Kentucky Snowberry Mountain unknown macrogroup & group Vegetation Herbaceous Semi-natural Wheatgrass Intermediate Shrubland Chaparral Sclerophyllous Interior American North Western - 2.B.2.Nd Chaparral Interior Cool - Ar patula Arctostaphylos Shrubland - Cliff-rose Manzanita Greenleaf Stansbury - Mountain Big Manzanita Greenleaf Sagebrus Manzanita ShrublandGreenleaf B1b. Chaparral Interior Warm A1b. Mesa gr - A1c. Up Ceanothus turbinella Quercus Shrubland Manzanita Mexican - Serviceberry) Live Colluvial (Utah Oak Turbinella A1c. Upland A1c. Upland A1a. Mesa 2.B.6.Nb 2.B.6.Nb Freshwat -North American Western Shrubland & Prairie Mixedgrass Plains Great curtipe - Bouteloua scoparium Schizachyrium Vegeta Herbaceous Semi-natural Wheatgrass) Smooth - Brome (Western Meadow Wet & Shrubland Wet Subalpine & Montane American North Western Group Wet Meadow Montane spp. - Calamagrostis spp. Carex SedgeBeaked Herbaceous Vegetation Herbaceous Rush Baltic Vegetation Sedge Herbaceous Nebraska Group Shrubland Seep spp. Riparian & Salix Willow / BeakedStrapleaf Sedg Semi-natural Bluegrass Kentucky B1a. High Platea / Hi Streambank B2a. 94 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8 lier) / Shrubland / Herbaceous / Sparse / Shrubland / Herbaceous / Shrubland Group.  es / Upland Sands (out es odland / Shrubland / Sparse / Sparse / Shrubland odland ed on coalition group affinities rest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland rest Forest / Woodland / Shrubland / Herbaceous / Shrubland / Woodland Forest ds - Woodland / Shrubland / Sparse / Sparse / Shrubland -ds Woodland Macrogroup ands / Lowland Flats -ands / Lowland Flats Woodland  bank - Forest / Woodland - Forest bank Lowlands - Woodland / Shrubland / Sparse / Sparse / Shrubland - Woodland Lowlands ands / Rocky Slopes / Upland (outlier) Sands Slopes / Rocky ands id Lowlands - Woodland / Shrubland / Sparse / Sparse / Shrubland - Woodland Lowlands id Lowlands / Rocky Slop Lowlands / Rocky c. Upland Sands / Rocky Slopes - / Sparse Slopes Shrubland / Rocky Sands c. Upland A2c. Upland Sands / Rocky Slopes - Shrubland / Sparse - Slopes Shrubland / Rocky Sands A2c. Upland X.X.X.Xx - Division  bland A2b. Arid Lowlands - Wo - Lowlands Arid bland A2b. X.X.X - Formation X.X.X - Formation  can Warm Desert Riparian Low Bosque & Shrubland Group Shrubland & Low Bosque Riparian Desert Warm can ens Rocky Mountain & Great Basin Riparian & Seep Shrubland Group Group Shrubland & Seep Basin Riparian & Great Mountain Rocky ens arex obnupta Vancouverian Freshwater Coastal Marsh & Meadow Group Group Meadow & Marsh Coastal Freshwater Vancouverian obnupta arex -natural Shrub Herbaceous Vegetation A2. - Ephedra torreyana Colorado Plateau Shrubland Group Group Shrubland Plateau Colorado torreyana - Ephedra Herbaceous VegetationHerbaceous - Fo Streambank B2b. X.X Subgrass  can Warm Desert Freshwater Marsh Marsh Desert can Warm Freshwater Shrubland A2a. Arid Lowl Arid Shrubland A2a. 7. Continued. X - Class PPENDIX A Association class Alternative bas Western North American Temperate Lowland Wet Shrubland, Wet Meadow & Marsh Marsh & Meadow Wet Shrubland, Wet Lowland Temperate American North Western - C egedii - Argentina beringensis Deschampsia Scouringrush) Variegated Horsetail, (Field pubesc Forestiera - spp. - exigua Crataegus Salix Shrubland Graminoids / Mesic Willow Coyote Ameri North Southwestern -2.B.6.Nc & Marsh Meadow Shrubland, Freshwater Desert Warm Ameri North spp. - Baccharis glandulosa Prosopis Arrow-weed Seasonally Flooded & Shrubland Meadow Wet Marsh, Freshwater Boreal & - Temperate 2.B.6 Shrubland / Barren Willow Coyote Shrubland Seepwillow Emory Semi-Desert & 3 - Desert & Grassland Scrub Semi-Desert - Cool 3.B & Grassland Scrub Semi-Desert -3.B.1 Cool Grassland & Scrub Semi-Desert Cool American North - Western 3.B.1.Ne Stream B2b. Grassland & Shrubland Dry Basin & Intermountain Great Group Shrubland lanata - Krascheninnikovia nauseosa - Ericameria viscidiflorus Chrysothamnus Vegetation Sparse Deposit Sand Rabbitbrush Rubber / Kentucky Bluegrass SemiRabbitbrush Green Semi-natural Shrubland / Cheatgrass Rabbitbrush Rubber Dwarf-shru / James' Galleta species) - (Prickly-pear Snakeweed A2. Lowl - Streambank B2b. A2 Coleogyne ramosissima - Ephedra cutleri cutleri Ephedra - ramosissima Coleogyne Shrubland / Galleta James' Blackbrush Shrubland Blackbrush Shrubland Plateau Colorado Sagebrush Sand A2b. Arid A2b. Ar Lowlan A2b. Arid 2015] PLANT COALITION GROUPS OF ZION NP 95

ceous / Sparse aceous / Sparse rubland / Herbaceous / Sparserubland bland / Herbaceous / Sparse rubland / Herbaceousrubland / Sparse rubland / Herbaceousrubland / Sparse - Woodland / Shrubland / Herba / Shrubland - Woodland es - Woodland / Shrubland / Herb land / Shrubland / Sparse / Sparse / Shrubland land cky - Slopes / Sh Woodland Group. & Steppe Group Group Steppe &  Flats -Flats / Shrubland Woodland / Herbaceous / Sparse d Flats -d Flats / Shrubland Woodland / Herbaceous / Sparse and Flats -and Flats / Shru Woodland ed on coalition group affinities wland Flats -wland Flats / Sh Woodland wland Flats -wland Flats / Sh Woodland Woodland / Shrubland / Sparse / Sparse / Shrubland Woodland rest / Woodland / Shrubland / Shrubland / Woodland rest Macrogroup  Sands / Slickrock / Rocky Slopes / Rocky Slopes / Slickrock Sands wlands / Lowland Top / Rocky Slopes - Woodland / Shrubland / Sparse / Sparse / Shrubland - Slopes Woodland Top / Rocky Lowlands / Lowlan nd Sands / Slickrock Rocky Slop nd Sands / Rocky Slopes - Shrubland / Sparse - Slopes Shrubland / Rocky nd Sands Lowlands / Lowl sa Top / Rocky Slopes - Slopes Wood sa Top / Rocky sa Top / Upland Sands / Slickrock / Rocky Slopes (outlier) / Rocky Slopes / Slickrock Sands / Upland sa Top Arid Lowlands / Lo Upland Sands / Rocky Slopes - Shrubland / Sparse - Slopes Shrubland / Rocky Sands Upland Upland Sands / Slickrock Ro

X.X.X.Xx - Division  Shrubland A2b. Arid Lowlands - Lowlands Arid Shrubland A2b. ye) Shrubland Fo Top - A1a. Mesa X.X.X - Formation X.X.X - Formation  innikovia lanata Scrub Group Group Scrub lanata innikovia tridentata ssp. vaseyana - Artemisia cana ssp. viscidula Tall Shrubland Tall viscidula cana ssp. - Artemisia vaseyana ssp. tridentata misia tridentata ssp. tridentata Tall Sagebrush Group Group Sagebrush Tall ssp. tridentata tridentata misia s - Hesperostipa comata Semi-Desert Grassland Group Group Grassland Semi-Desert comata s - Hesperostipa - Purshia tridentata Big Sagebrush Steppe Group Group Steppe Sagebrush Big tridentata - Purshia X.X Subgrass esert Cliff, Scree & Rock Vegetation Vegetation Rock Scree & Cliff, esert  ush Shrubland A2a. Arid ead Herbaceous Vegetationead Herbaceous A1c. Upla land A2a. Arid Lowlands / Lo Herbaceous Vegetation A2a. Arid Lo a - Atriplex canescens - Kraschen canescens a - Atriplex 7. Continued. X - Class PPENDIX A Association class Alternative bas 6 - Rock Vegetation Vegetation 6 - Rock Rock Vegetation Semi-Desert 6.C - & Desert Rock Vegetation & Scree Cliff, Semi-Desert Warm -6.C.1 Semi-D Warm American 6.C.1.Na - North Rock Vegetation Scree & Cliff, Semi-Desert Warm North American Nevada Joint-fir Basalt Shrub Pleuraphis jamesii - Achnatherum hymenoide - Achnatherum jamesii Pleuraphis Blue Grama - Needle-and-Thr Herbaceous VegetationJames' Galleta Vegetation Herbaceous Basin Needle-and-Thread Great Sand Dropseed Great Basin Steppe & Shrubland Sage Dwarf Basin & Intermountain Great Group & Steppe Shrubland nova - Artemisia bigelovii Artemisia - arbuscula Artemisia Dwarf-shrubland / Bottlebrush Sagebrush Black Dwarf-shrubland / Muttongrass Sagebrush Black Sagebrush / Needle-and-Thread Dwarf-shrubland Black Steppe & Shrubland Sagebrush Tall Basin & Intermountain Great tripartita Artemisia - tridentata Artemisia A1. Me R Wild - (Streamside Wheatgrass / Western Sagebrush Big Basin Artemisia - spiciformis ssp. tridentata Artemisia / Needle-and-Thread Sagebrush Big Mountain Shrubland A2a. Arte - wyomingensis ssp. tridentata Artemisia Semi-natural / Cheatgrass Rabbitbrush) - Sagebrush Big (Rubber Big Grama Sagebrush/ Blue Shrubland A2c. A1c. Upland Scrub Saltbrush Great Basin A1b. Mesa confertifoli Atriplex -Fourwing Saltbush Big Sagebr Fourwing Saltbush Shrubland - Mormon-tea Shrubland Horsebrush Gray A1b. Me A1c. A2c. Upla A2a. Arid 96 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [Volume 8

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nd / Shrubland / Sparse / Sparse nd / Shrubland ed on coalition group affinities ds / Lowland Flats -ds / Lowland Flats Woodland Macrogroup  sa Top / Rocky Slopes - Slopes Wood sa Top / Rocky A2b. Arid Lowlands - Woodla Lowlands A2b. Arid A1c. Upland Sands / Slickrock Rocky Slop r Forest and Woodland 1 X.X.X.Xx - Division  Sparse Vegetation Group Vegetation Sparse Ravine WoodlandRavine Mesic Montane Mixed Conife Mixed Montane Mesic X.X.X - Formation X.X.X - Formation sic Montane Mixed Conifer Forest and Woodland Forest Conifer Mixed sic Montane 1  Active and Stabilized Dune and Woodland Forest Conifer Aspen-Mixed a spp. Intermountain Basins Intermountain a spp. Bigtooth Maple Basins Shale Badland ert Cliff, Scree & Rock Vegetation Rock Vegetation Scree & ert Cliff, in Aspen Forest and Woodland X.X Subgrass  se Vegetation A2a. Arid Lowlan 7. Continued. X - Class 8. Numbers of associations of the U.S. National Vegetation Classification (NVC) at Zion National Park, Utah (Cogan et al. 2004) Classification (NVC) at Zion National Park, National Vegetation 8. Numbers of associations the U.S. PPENDIX PPENDIX Land Cover Class Ecological System A1a A1b A1c A2a A2b A2c A2d B1a B1b B2a B2b A Association class Alternative bas A 6.C.2 - Cool Semi-Desert Cliff, Scree & Other Rock Vegetation Vegetation & Other Rock Scree Cliff, Semi-Desert 6.C.2 - Cool Semi-Des Cool American 6.C.2.Na - North Vegetation Sparse Badland Scree & Cliff, Basins Intermountain - Ephedr spp. - spp. Cercocarpus Atriplex Vegetation Sparse Pavement Mountain-mahogany Rock Vegetation Sparse Badlands Buckwheat Wild Crispleaf Nevada Joint-fir Spar / Lichen Ponderosa SlickrockVegetation Pine Sparse A1b. Me Barren Inter-Mountain Basins Barren Barren Inter-Mountain Barren Barren Inter-Mountain Barren and Tableland Canyon Mixed Bedrock Plateau Colorado Woodland and Forest Woodland Pinyon-Juniper Plateau Colorado Woodland and Forest Basins and Canyon Cliff Inter-Mountain Woodland Pinyon-Juniper Basin Great Woodland and Forest Basins Inter-Mountain Forest and Woodland Basins Rock Volcanic Inter-Mountain Land and Cinder Rocky Mounta Forest and Woodland Rocky Mountain Woodland and Forest Dry- Mountain Rocky Southern Woodland and Forest Me Mountain Rocky Southern 1 4 2 logical systems (obtained or inferred from NatureServe 2013) and alternative classes based on coalition group affinities (see F 2015] PLANT COALITION GROUPS OF ZION NP 97

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odland and Shrublandodland oodland and Shrubland wer Montane Riparian Wo Riparian wer Montane e-Foothill Riparian Woodland and Shrubland ane-FoothillW Riparian rt Riparian and Shrubland Woodland rt lpine-MontaneWoodland Riparian untain Ponderosa Pine Woodlanduntain Ponderosa Pine 2 1 Alpine-Montane Wet Meadow 8. Continued. PPENDIX Land Cover Class Ecological System A1a A1b A1c A2a A2b A2c A2d B1a B1b B2a B2b A Herbaceous Inter-Mountain Basins Semi-Desert Grassland Semi-Desert Basins Woodland and Forest Mo Rocky Southern Herbaceous Inter-Mountain Herbaceous Wetland Rocky Mountain Herbaceous Wetland Rocky Mountain Lower Mont Shrubland Shrubland Shrubland Shrubland Blackbrush-Mormon-Tea Shrubland Plateau Colorado Shrubland Sagebrush Low Shrubland Mixed Plateau Colorado Shrubland Shrubland Pinyon-Juniper Plateau Colorado Shrubland Shrubland Sagebrush Mixed Xeric Basin Great Shrubland Basins Big Shrubland Sagebrush Inter-Mountain Chaparral Shrubland Desert Scrub Salt Basins Mixed Inter-Mountain Mogollon Shrubland Steppe/Savanna Oak-Mixed Shrubland Gambel Montane Mountain Rocky Steppe/Savanna Shrubland Montane-Foothill Lower Mountain Rocky Steppe Sagebrush Basins Big Inter-Mountain Steppe/Savanna Sand Shrubland Plateau Colorado Southern Steppe Sagebrush Basins Montane Inter-Mountain Wetland Woody Shrub-Steppe Basins Semi-Desert Inter-Mountain Wetland Woody Lo Warm Desert American North Wetland Woody 1 Warm Dese American North Wetland Woody 3 1 Montan Lower Mountain Rocky Woody Wetland 1 Subalpine-Montane Mountain Shrubland Rocky Riparian Rocky Mountain Suba 1 1 1 2