MADRONÄ O, Vol. 52, No. 4, pp. 222±257, 2005

SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA: A DATABASE OF SERPENTINE AFFINITY

H. D. SAFFORD1,2,J.H.VIERS3, AND S. P. HARRISON2 1 USDA-Forest Service, Paci®c Southwest Region, 1323 Club Drive, Vallejo, CA 94592 [email protected] 2 Department of Environmental Science and Policy, University of California, Davis, CA 95616 3 Information Center for the Environment, DESP, University of California, Davis, CA 95616

ABSTRACT We present a summary of a database documenting levels of af®nity to ultrama®c (``serpentine'') sub- strates for taxa in the California ¯ora, USA. We constructed our database through an extensive literature search, expert opinion, ®eld observations, and intensive use of accession records at key herbaria. We developed a semi-quantitative methodology for determining levels of serpentine af®nity (strictly endemic, broadly endemic, strong ``indicator'', etc.) in the California ¯ora. In this contribution, we provide a list of taxa having high af®nity to ultrama®c/serpentine substrates in California, and present information on rarity, geographic distribution, taxonomy, and lifeform. Of species endemic to California, 12.5% are restricted to ultrama®c substrates. Most of these taxa come from a half-dozen plant families, and from only one or two genera within each family. The North Coast and Klamath Ranges support more serpentine endemics than the rest of the State combined. 15% of all plant taxa listed as threatened or endangered in California show some degree of association with ultrama®c substrates. Information in our database should prove valuable to efforts in ecology, ¯oristics, biosystematics, conservation, and land management. Key Words: serpentine, ultrama®c, California, endemism, diversity.

INTRODUCTION lain by ultrama®c rocks (6000 km2/406,280 km2). Ultrama®c rocks, often called ``serpentine'' by In addition, because they tend to have small geo- ecologists, botanists and pedologists, underlie more graphic ranges and because many of them occur in than 6000 km2 of the land area of the State of Cal- the rapidly urbanizing San Francisco Bay Area, ser- ifornia (Harrison et al. 2000). The edges of conti- pentine endemics are overrepresented among the nental plates often include bands of these vestiges state's rare, sensitive, and listed plant taxa (Skinner of oceanic mantle rock, accreted during the geolog- and Pavlik 1994). The ecology of California's ser- ic process of subduction, and later uplifted and ex- pentine plants has been extensively studied at the posed during mountain building and subsequent University of California's Sedgwick Ranch Reserve erosion. Ultrama®c rocks and the soils that develop (e.g., Seabloom et al. 2003; Gram et al. 2004) and on them are characterized by critically low levels McLaughlin Reserve (e.g., Harrison et al. 2003; of most principal plant nutrients (N, P, K, Ca), and Safford and Harrison 2004) and Stanford Univer- exceptionally high levels of Mg and Fe and a suite sity's Jasper Ridge Reserve (e.g., McNaughton of toxic trace elements including Cr, Ni, and Co. 1968; Huenneke et al. 1990; Hobbs and Mooney Outcrops of ultrama®c rocks support high numbers 1991). of edaphic-endemic taxa throughout the world Botanists have relied for two decades on the (Brooks 1987). The California serpentine ¯ora is monograph by Arthur Kruckeberg (1984) for most the richest in the temperate zone, and consists of of their information on Californian serpentine-en- hundreds of species and subspecies that are largely demic plant taxa. Since then, publication of the Jep- or entirely con®ned to ultrama®c substrates. son Manual (Hickman 1993), and a proliferation of Serpentine endemism is a key feature of the di- new botanical research and name changes have left versity of the California ¯ora (Raven and Axelrod this list in need of updating. Our initial aim was to 1978; Kruckeberg 2002). Of about 1410 full spe- modify Kruckeberg's (1984) list, primarily using cies endemic to the State (Hickman 1993), Kruck- information from Hickman (1993), to use in our eberg (1984) estimated that about 180 were endem- research on diversity patterns (Harrison et al. 2000, ic to serpentine. If these numbers are at least ap- 2004). However, it soon became clear that we proximately correct, then about 13% of the plant would have to expand and intensify our search for species endemic to California are serpentine en- the best available information. Complicating this demics. This is a remarkably high number when effort, plants show a continuum in degrees of ser- one considers that only 1.5% of the State is under- pentine restriction, and are sometimes more restrict- 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 223 ed in some parts of their geographic ranges than marks (``?'') were attached to those taxa for which others, thus contributing to inconsistencies among more information was necessary to con®dently as- reports from different sources. This led us to adopt sign their status. Some of the ``tentative'' endemics a semi-quantitative procedure for scoring plant taxa were included in the indicator appendix as well, on their reported degree of serpentine af®nity. thus these taxa occur twice in Kruckeberg's lists. In this contribution, we present a summary of our We combined Kruckeberg's two scales, and added current database of serpentine af®nity in the Cali- two levels to yield six levels of ultrama®c af®nity, fornia ¯ora. The database was constructed via an where 6 represents a ``strict endemic'' (Ն95% of extensive literature search, expert opinion, ®eld ob- occurrences on ultrama®cs), and successively lower servations, web research, and intensive use of ac- values signify lower af®nity to the substrate (5 ϭ cession records at key herbaria. It provides data on 85±94% of occurrences; 4 ϭ 75±84%; 3 ϭ 65± levels of serpentine endemism, rarity, geographic 74%; 2 ϭ 55±64%; 1 ϭ 45±54%). By this de®ni- distribution, taxonomy, and lifeform. tion, ``1'' thus represents a species found about half of the time on serpentine. We consider scores be- METHODOLOGY tween 1 and 2 to indicate ``weak indicators'', and a score of about 1 to mean an ``indifferent'' taxon. We began by conducting a database search of the The Kruckeberg ®delity scale crosswalks to ours in electronic Jepson Manual (Hickman 1993) main- the following fashion: ``ϩϩϩ'' ϭ 6; ``ϩϩ'' ϭ 5; tained by the Jepson Herbarium at the University ``!!'' ϭ 3; one ``!'' ϭ 2. Those taxa which occurred of California-Berkeley (UC-JEPS 2004a). The da- in both Kruckeberg's endemic and indicator tables tabase was queried for all taxa with ``serpentine'', had their two scores averaged: these all fell be- ``ultrama®c'', or related (e.g., ``asbestos soils'') ref- tween ``3'' and ``4'' on our scale. For example, Cu- erences in the habitat description. Taxa containing pressus macnabiana was rated ``ϩϩϩ'' in Kruck- ``non-serpentine'' in the description were removed eberg's Appendix C (i.e., ``6'' on our scale), and afterward. We cross-checked the 391 serpentine-re- ``!!'' [i.e., ``2'' in our scale] in Appendix D; these lated taxa found in the Jepson Manual with Kruck- were averaged to ``4'' on our scale. eberg (1984), who listed those taxa he believed to We attached our categorical levels of ultrama®c be endemic to ultrama®c substrates in California, af®nity to all of the species in our hybrid Jepson- and those that were either local or regional ``ser- Kruckeberg database. In the case of the Kruckeberg pentine indicators'' (i.e., nonendemic taxa whose taxa, we simply cross-walked the Kruckeberg ®- distributions are nonetheless skewed toward occur- delity codes to our scale as described above, mak- rences on ultrama®cs). Taxonomic updates in the ing some adjustments based on more recent taxo- Jepson Manual (Hickman 1993) were applied to the nomic revisions and combinations. In the case of Kruckeberg list (which included 377 taxa after the Jepson Manual taxa, we were forced to interpret these revisions), and then those taxa not on the Jep- the language used in habitat descriptions to deter- son-derived list were added to our database. This mine levels of af®nity. We used the following in- resulted in a list of 529 taxa; of these, 287 were not terpretations of description language to assign af- shared between the two sources. We then added to ®nities: a ``6'' was assigned where the habitat de- the list a number of taxa that we considered to be scription categorically stated ``serpentine'' or ``ul- likely endemics or indicators but which were not trama®c'' (a ``5'' if there was some indication that indicated as such by either Kruckeberg (1984) or this restriction was not absolute); a ``4'' was as- the Jepson Manual (1993). Finally, published lit- signed where the modi®ers ``generally'' or ``usually erature (e.g., Meinke and Zika 1992; Nelson and serpentine'' were used; ``especially'' or ``often'' Nelson 2004; Baldwin 1999 and 2001; Barkley equaled ``3''; ``sometimes'' or ``occasionally'' 1999; Porter and Johnson 2000; Zika et al. 1998) equaled ``1''. In a few cases, af®nity levels were and the online Jepson Interchange Jepson Flora assigned based on ancillary information in the hab- Project (UC-JEPS 2004b) were consulted for tax- itat and/or range description rather than on explicit onomic revisions and taxa newly described since statement of serpentine af®nity. the publication of the Jepson Manual. We then conducted a broad survey of the litera- To score the af®nity of taxa to ultrama®c sub- ture, regional botanical experts, and herbaria rec- strates, we adopted a modi®cation of Kruckeberg's ords to obtain as many sources as possible for each measures of ultrama®c ``®delity''. In his Appendix taxon in our database, and to add to the database C, Kruckeberg (1984) used two or three ``ϩ''s to any taxa we might have overlooked. We manually signify increasing levels of endemism: three ``ϩ''s consulted every species description in a variety of were attached to taxa with 95±100% of their oc- regional and local ¯oras (Clifton 2001; Ertter and currences found on ultrama®cs, two ``ϩ''s signi®ed Bowerman 2004; Howell 1970; McMinn 1939; Os- taxa with 85±94% ®delity. In his Appendix D, wald 2002; Smith and Wheeler 1992), and guide- Kruckeberg used one or two exclamation marks books to rare and sensitive taxa (Hanson 1999; (``!''s) to signify increasing levels of ®delity to ul- Hoover et al. 1993; Jimerson et al. 1995; McCarten trama®c substrates among supposed nonendemic 1988; McCarten and Rogers 1991; Nakamura and ``indicator'' taxa. In both appendices, question Nelson 2001; Trinity SIPS 2001; USFWS 1998). 224 MADRONÄ O [Vol. 52

We also consulted the CalFlora Online Species Da- In our online accession database research, we tabase (CalFlora 2004), and the California Native followed the following protocol: Plant Society Online Inventory of Rare and Endan- 1. We began with the most recent accession re- gered Plants (CNPS 2004). We added columns to cords and worked backwards, as habitat descrip- our database for each source, and gave scores (1± tions before the mid 1970's usually lack suf®ciently 6, as described above) to each taxon for which a detailed information on substrate and location. habitat description suggested an ultrama®c af®nity. 2. We consulted the habitat description for each Information on serpentine af®nity in the CalFlora record. If the description included enough infor- database is limited to taxa from the Sierra Nevada mation to determine the substrate, we noted wheth- and to rare taxa statewide, and does not include er it was ultrama®c or non-ultrama®c. We did not suf®cient information to determine degree of af®n- count multiple accession records from the same ity (A. Dennis, personal communication). CalFlora collecting trip and location as different records. was therefore not treated as a typical ``source'', and 3. On the average, about ⅓ of the accession rec- CalFlora serpentine taxa were simply given a score ords consulted had suf®cient information to deter- of 0.5 to be added later to the sum of scores when mine if a collection had been made on ultrama®cs ®nal ultrama®c af®nities were calculated (see be- or not. Not all of these determinations were made low). The California Natural Diversity Database simply based on the collector's habitat description. (CNDDB) was not searched, as we consulted all of For example, many California counties do not con- the primary resources originally used to build tain outcrops of ultrama®c rocks (e.g., Los Angeles, CNDDB, and the CNPS Online Inventory (see San Diego, San Bernardino, Modoc). Collections above) is updated from the same contemporary from these counties were coded as ``nonserpentine'' sources as CNDDB. even where habitat descriptions were missing. Also, We calculated preliminary mean af®nities for collections from well-known collecting locations on taxa in our database by summing the scores across ultrama®cs (e.g., Blue Banks in Glenn County, Red source columns and adding the CalFlora score (if Butte in Siskiyou County, or the mouth of 18-Mile present), then dividing by the number of sources Creek on the Middle Fork Smith River, Del Norte (not including CalFlora) for the taxon in question. County) were coded as ``serpentine'' even where We also calculated the number of sources, the me- habitat descriptions were missing. Finally, where dian score, and the standard deviation and standard we had trouble getting a suf®cient number of rec- error of the scores for each taxon. We then sent the ords with habitat descriptions, or where it was oth- database to approximately 40 state and regional ex- erwise critical to get more information, we used perts for their review and input, and asked them to location information in the accession record (where score serpentine af®nity using the 1±6 scale for it existed) to do further research. We used TOPO! taxa with which they were familiar. These individ- Software (National Geographic Maps 2000) to lo- uals included botanists employed by federal and cate coordinates or named locations and then con- state land management agencies, universities, mu- sulted geological maps (ranging from 1:250,000 to seums, non-governmental organizations, and pri- 1:25,000) to determine if the location was on an vate consulting ®rms. We received 17 substantive ultrama®c outcrop. Only those occurrences which replies, and incorporated their input into an updated could be con®dently assigned to ultrama®cs were database. identi®ed as such. The next step was to ensure that we had at least 4. We continued until we had recorded habitat three sources of serpentine af®nity for each taxon information from at least 10% of the total accession in our database; given the great differences between records for the species in question. Our minimum the Jepson Manual and Kruckeberg's list, we felt a was 10 records, unless there were fewer than 10 third opinion was important. We focused on those records with habitat descriptions and reasonably lo- taxa for which we had less than three sources, as catable site information (286/548 taxa had fewer well as those with high variability in scores. We than 10). Our maximum was usually 20, although began by consulting the habitat descriptions in we went beyond 20 in some cases. Munz and Keck (1968) for every taxon in our da- 5. We summarized the accession record results tabase with less than three sources. We then turned for each taxon by dividing the total number of rec- to Herbaria accession records. We searched the on- ords with suf®cient habitat or location information line ``SMASCH'' accession databases of the UC to determine substrate by the number of records and Jepson Herbariums at UC-Berkeley at (UC- recording serpentine/ultrama®cs, and then multi- JEPS 2004c) for all taxa with one or two sources, plied the result by 100 to get a percentage. We then and for all taxa with af®nity-score standard devia- cross-walked the percent value to our scale of ul- tions Ն 1.0 (a total of 548 taxa). For any Northern trama®c af®nity: 95±100% of records on ultramaf- California taxa remaining with less than three ics ϭ 6; 85±94% ϭ 5; 75±84% ϭ 4; 65±74% ϭ 3; sources and/or high variability, we then searched 55±64% ϭ 2; 45±54% ϭ 1; 35±44% ϭ 0.75; 25± the online accession database of the Biological Sci- 34% ϭ 0.5; 15±24% ϭ 0.25; Ͼ0±14% ϭ 0.1; 0 ences Herbarium at Chico State University (CSU- ϭ 0. BSH 2004; a total of 164 taxa were searched). Finally, T. Nelson and S. Carothers also used the 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 225

Humboldt State University Herbarium to provide and seven sources for 19 taxa; 587 taxa had be- information to us on a number of under-document- tween three and six sources. Eighty-one taxa had ed taxa from Northwestern California. fewer than three sources (77 with two, three with In our accession records research, we necessarily one). Somewhat more than half of the taxa (387) assumed that: (1) the taxon itself was correctly in our original list had standard deviations for ser- identi®ed on the accession record; (2) the substrate pentine af®nities Ͼ 1.0. was correctly identi®ed by the collector; and (3) Since our serpentine af®nities are calculated as ultrama®c substrates were neither more nor less the means of multiple sources, our values fall on a likely to be identi®ed correctly (or at all) than other continuous scale, rather than in categories. Given substrates. The last assumption is probably ¯awed, this, we recognized taxa with mean af®nities Ͼ 5.5 as serpentine and other ``charismatic'' substratesÐ as ``strict endemics'' (analogous to Kruckeberg's given their close connection to plant endemic taxa ``ϩϩϩ'', or taxa with Ͼ 95% of their occurrences and their relative ease of identi®cationÐare almost on ultrama®cs), and taxa with mean af®nities Ͼ 4.5 certainly more likely to be identi®ed than ``normal'' and Ͻ 5.5 as ``broad endemics'' (analogous to substrates. This could theoretically lead to acces- Kruckeberg's ``ϩϩ'', taxa with about 85±94% of sion records ``overstating'' the degree of a taxon's their occurrences on ultrama®cs). Using these def- af®nity to ultrama®c substrates. In practice, how- initions, 164 taxa are strict endemics, while 82 taxa ever, we found that the accession records were gen- are broad endemics, for a total of 246 endemic taxa; erally somewhat more conservative than our liter- 176 of these are full species. Among the remaining ature sources vis-aÁ-vis the serpentine af®nities of taxa, 123 are ``strong serpentine indicators'' the taxa in our database. (Kruckeberg 1984), with scores ranging from 2.5 Our ®nal database included 18 columns of infor- to 3.4 (about 65±74% of their occurrences on ul- mation sources for serpentine af®nity, plus a col- trama®cs); 150 are ``weak indicators'', falling be- umn for CalFlora. We summed these af®nity values tween 1.5 and 2.4 on our scale (Ϯ 55±64% of their and took their mean (not including CalFlora in the occurrences on ultrama®cs); and 79 fall in a gray denominator). We also calculated the mean without area between weak indicators and indifferent taxa CalFlora, the median, the standard deviation, and (between 1.0 and 1.4 on our scale, or about 50± the standard error. We identi®ed each taxon by tax- 54% of occurrences. Seventy-one taxa have af®nity onomic category (pteridophyte, gymnosperm, dicot scores between 3.5 and 4.4 (about 75±84% of their or monocot), and by lifeform (annual forb, peren- occurrences on ultrama®cs), and thus represent the nial forb, annual graminoid, perennial graminoid, transition from strong indicators to broad endemics. shrub, tree). For rare taxa, we added the rarity rat- Six families account for more than half of all the ing from the California Native Plant Society Online endemics: Asteraceae, Liliaceae, Brassicaceae, Po- Database of Rare and Endangered Plants (version lygonaceae, Scrophulariaceae, and Apiaceae (Table 6.04d, 11-12-2004). The following information was 1). The 20 most important plant families among the also added to the complete database: geographic serpentine endemics are shown in Fig. 1, with the distribution in California for each taxon (by Jepson percentage of the serpentine endemic ¯ora that they Manual geographic subdivisions); elevational range contribute, as well as the percentage of the total (from Hickman 1993); the geographic distribution California endemic ¯ora that they contribute. Fam- of, and number of species of the genus of each ilies that proportionally contribute more to the ser- taxon (from Mabberly 1996); and the common pentine endemic ¯ora than to the California endem- name (from Hickman 1993, and the Natural Re- ic ¯ora include Liliaceae, Brassicaceae, Polygona- source Conservation Service PLANTS online da- ceae, Linaceae and Caryophyllaceae. Families tabase [USDA-NRCS 2005]). Aside from a sum- whose level of endemism is much lower on serpen- mary of the geographic distribution, this informa- tine than it is statewide include Fabaceae, Poaceae, tion is not presented in the current paper, but is Boraginaceae, and Rosaceae (Fig. 1). available on request from the ®rst author, as are the The most diverse genera in our list of serpentine af®nity values calculated for each source. endemics are Streptanthus (Brassicaceae) and Er- iogonum (Polygonaceae), followed by Hesperoli- RESULTS non (Linaceae) and Arctostaphylos (Ericaceae) (Ta- ble 2). There are 21 genera with at least four taxa A summary table of the current database is pre- among the endemics. These represent 14 plant fam- sented in Appendix 1. Appendix 1 includes 669 ilies, with Asteraceae (four genera among the en- taxa, ranging in af®nity from 6.25 to 1.00 (some demics), Liliaceae (three genera), Scrophulariaceae values exceed 6 because they were identi®ed as ser- (two genera) and Brassicaceae (two genera) the pentine taxa in the CalFlora Database). Our full da- only families with multiple genera in the list. Figure tabase includes 698 taxa, 29 of which have mean 2 compares the contribution of these genera to the serpentine af®nities of Ͻ 1; we did not include serpentine endemic ¯ora with their contribution to these taxa in the current paper. The greatest number the California endemic ¯ora. All but ®ve or six of of sources we located for any single taxon was nine these genera have a greater level of endemism to (four taxa). We found eight sources for eight taxa serpentine than they have within the State as a 226 MADRONÄ O [Vol. 52

TABLE 1. NUMBERS OF SERPENTINE ENDEMIC AND NEAR tostaphylos contributes a similar percentage to both ENDEMIC TAXA, BY FAMILY. 1 Strict endemics. 2 Strict en- ¯oras (Fig. 2). demics plus broad endemics. 3 Strict and broad endemics Of the taxa in our database, there are 532 dicots plus ``near endemic'' taxa (taxa transitional from strong (of which 204 are endemic), 119 monocots (38 en- indicators to broad endemics). demics), 12 gymnosperms (2 endemics) and six Serpentine af®nity score pteridophytes (2 endemics). 207 taxa are annual Total forbs (of which 71 are endemics, including 7 of 14 Ն 1 Ն 2 Ն 3 Family 5.5 4.5 3.5 taxa that can also be perennial/biennial), 383 are peren- Asteraceae 26 37 45 106 nial forbs (150 endemics, including the 7 ``annu- Liliaceae 15 28 37 85 als'' and 6 taxa which can also be shrubs), 24 are Brassicaceae 21 26 31 46 perennial graminoids (7 endemics), 64 are shrubs Polygonaceae 10 17 19 39 (23 endemics, including 6 taxa shared with the pe- Scrophulariaceae 9 14 18 37 rennial forbs and 1 which assumes both tree and Apiaceae 7 10 13 32 Linaceae 8 9 9 14 shrub forms), and 12 are trees (2 endemics) (Ap- Ericaceae 5 8 10 15 pendix 1). Of the endemic perennial forbs, 24 are Polemoniaceae 6 7 8 18 bulb plants (all Liliaceae), 17 are rhizomatous Caryophyllaceae 5 7 8 18 (from ten different Families), three are hemipara- Fabaceae 4 7 10 24 sites (Scrophulariaceae), and one is carnivorous Lamiaceae 4 6 10 17 (Lentibulariaceae) (Appendix 1). Crassulaceae 5 5 7 13 Using Kruckeberg's (1984) physiographic prov- Rhamnaceae 4 5 6 14 inces of California (which correspond more or less Campanulaceae 3 5 8 12 to major geographic subdivisions mapped in the Onagraceae 3 5 7 12 Hydrophyllaceae 4 4 8 15 Jepson Manual (Hickman 1993)), we found the fol- Rubiaceae 3448lowing geographic distribution of serpentine en- Convolvulaceae 1456demic taxa (Fig. 3): The North Coast, considered Cyperaceae 1458in toto (i.e., the Jepson Manual's NCo and NCoR Poaceae 1 3 3 19 subregions (Hickman 1993)), supports approxi- Portulacaceae 0 3 5 16 mately 118 serpentine endemics, with 49 of these Boraginaceae 2 2 3 10 restricted to that area. The Klamath Region (Jepson Gentianaceae 2223Manual subregion KR), supports 98 endemic taxa, Iridaceae 2224with 54 restricted to that area (including taxa also Malvaceae 2225 Salicaceae 2223found in neighboring SW Oregon). The San Fran- Garryaceae 1222cisco Bay Area (Jepson Manual subregion SnFrB Rosaceae 1 2 5 10 plus the sections of NCo and CCo bordering it) Cupressaceae 0236supports about 51 endemics, with 24 found only Violaceae 0237there. The South Coast Ranges, including the Chan- Asclepiadaceae 1111nel Islands and the Santa Ana Mountains (i.e., Jep- Berberidaceae 1114son Manual subregions CCo, SCoR plus the few Dryopteridaceae 1112ultrama®c outcrops that occur in the Jepson SW Fagaceae 1113Region), support 43 total endemics with 24 restrict- Lentibulariaceae 1111 Papaveraceae 1115ed to that area. The Sierra Nevada (Jepson Manual Ranunculaceae 1136region SN) support 38 total serpentine endemic Orchidaceae 0113taxa, with 21 taxa restricted to the Range (Fig. 3). Pteridaceae 0114 Of the 669 taxa in our database, 295 are listed Verbenaceae 0111as ``rare'' or ``uncommon'' by the California Native Cistaceae 0001Plant Society (CNPS) (Appendix 1). These include Orobanchaceae 0001194 of the 246 taxa that we consider to be either Pinaceae 0016strict or broad serpentine endemics. One serpentine Plantaginaceae 0001 Polygalaceae 0001endemic taxon, Arctostaphylos hookeri subsp. fran- Primulaceae 0001ciscana, is extinct in the wild and survives only in Sarraceniaceae 0011cultivation. Of the 295 rare or uncommon taxa, 154 Saxifragaceae 0012are on CNPS List 1b, which lists plants considered Sterculiaceae 0001threatened or endangered by either the State or Fed- Totals 164 246 315 669 eral governments, as well as unlisted plants which CNPS considers rare enough to warrant listing; 111 of these List 1b plants are serpentine endemics by our de®nition. Nine taxa (seven endemics) from whole. These genera include Streptanthus, Hesper- Appendix 1 are on CNPS list 2, which contains olinon, Lomatium and Minuartia. Only one genus plant taxa that are rare in California but are not (Phacelia) contributes less to the serpentine endem- restricted completely to the State; all of these taxa ic ¯ora than it does to the State as a whole; Arc- are either State listed and threatened or endangered, 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 227

FIG. 1. The twenty most important plant families of serpentine endemic plants (i.e., including strict and broad ser- pentine endemics), with the percentage of endemic species they contribute to the serpentine endemic ¯ora in California, and to the California endemic ¯ora as a whole. or are eligible for listing. Eight taxa (four endem- DISCUSSION ics) in Appendix 1 are found on CNPS list 3, which lists uncommon taxa for which more information is In 1984, Kruckeberg estimated that the serpen- required. Of taxa in Appendix 1, 123 (71 endemics) tine endemic ¯ora of California numbered approx- are on CNPS list 4, which contains taxa of ``limited imately 220 taxa (about 180 full species), and that distribution or infrequent throughout a broader area a further 230 taxa were suf®ciently associated with in California''. ultrama®cs to be ``indicators'' of the substrate. Thus, Kruckeberg believed that about 450 taxa were associated with serpentine in California. Al- TABLE 2. GENERA WITH MORE THAN THREE TAXA ENDEMIC though our results suggest that the number of ser- TO SERPENTINE. pentine-associated taxa is closer to 670, they also Endemic suggest that Kruckeberg's (1984) estimate of the Genus Family taxa number of full-species endemics was remarkably accurate (180 vs. 176). As Kruckeberg's numbers Streptanthus Brassicaceae 18 also suggested, serpentine endemics therefore com- Eriogonum Polygonaceae 14 prise approximately 12.5% (176/1410) of the plant Hesperolinon Linaceae 9 Arctostaphylos Ericaceae 8 species endemic to California. Based on numbers Allium Liliaceae 7 from the Jepson Manual (Hickman 1993; R. Moe Lomatium Apiaceae 7 personal communication), the percentage of serpen- Packera (Senecio) Asteraceae 6 tine endemics among California endemic species, Calochortus Liliaceae 5 subspecies and varieties is about 11.4% (246/2153). Cordylanthus Scrophulariaceae 5 Kruckeberg's (1984) estimates of endemics by Arabis Brassicaceae 4 California geographic region are somewhat less ac- Calystegia Convolvulaceae 4 curate than his statewide estimate (see Fig. 3), but Carex Cyperaceae 4 Castilleja Scrophulariaceae 4 Kruckeberg's data sources in the 1970's and early Cirsium Asteraceae 4 1980's were extremely limited compared to ours. Erigeron Asteraceae 4 As did Kruckeberg, we found that the North Coast Fritillaria Liliaceae 4 Ranges support more serpentine endemics plants Galium Rubiaceae 4 than any other geographic region, but that the Lessingia Asteraceae 4 Klamath Ranges (and adjoining SW Oregon) sup- Minuartia Caryophyllaceae 4 port many more restricted endemics than Krucke- Monardella Lamiaceae 4 berg thought was the case (54 vs. 30). Kruckeberg's Phacelia Hydrophyllaceae 4 estimates for the numbers of restricted endemics in 228 MADRONÄ O [Vol. 52

FIG. 2. The twenty-one most important genera of serpentine endemic plants (i.e., including strict and broad serpentine endemics), with the percentage of endemic species they contribute to the serpentine endemic ¯ora in California, and to the California endemic ¯ora as a whole. the North Coast Ranges and the Bay Area are very toriously uninformative when it came to habitat de- similar to our numbers (Fig. 3), but he overesti- scription. In contrast, many data sources we ac- mated the number of endemics in the South Coast cessed were available electronically and could be Ranges (36 vs. 24). Kruckeberg estimated that ei- queried and retrieved remotely. ther 13 or 16 (depending on whether one goes by The other principal reason for difference is the the text or the tables in Appendix E) endemic taxa inevitable discoveries and taxonomic reorganiza- were restricted to the Sierra Nevada; we found 21 tions that occur over a 20-year period. Krucke- taxa thus restricted. berg's work came before publication of the Jepson Reasons for differences between our numbers Manual (Hickman 1993), which contained many and Kruckeberg's (1984) are many, but belong to signi®cant changes in California plant taxonomy. A two broad categories. The primary reason is quality considerable number of serpentine endemic taxa in and quantity of information. In many cases, Kruck- the Jepson Manual were wholly unknown to Kruck- eberg's information had to come through his own eberg in 1984. Examples include Calochortus ®eld experience, or through hard copy herbarium raichei S. Farwig & V. Girard, Minuartia stoloni- records, whichÐbefore the late 1970'sÐwere no- fera T. W. Nelson & J. P. Nelson, Perideridia ba- cigalupii Chuang & Constance, and Monardella stebbinsii Hardham & J. Bartel. Since the Manual's publication, there have been further changes (e.g., Barkley 1999; Baldwin 1999; Porter and Johnson 2000). Serpentine endemic taxa named since pub- lication of the Jepson Manual include Harmonia guggolziorum B. G. Baldwin, Carex serpentinicola P. F. Zika, and Silene serpentinicola T. W. Nelson & J. P. Nelson. As a null hypothesis, one might expect that the distribution of endemic plant taxa across plant fam- ilies and genera on California serpentines would more or less mirror the distribution of endemics in FIG. 3. Geographic distribution of serpentine endemic the State as a whole. Our data demonstrate that this taxa in California. ``Total endemics'' includes all Califor- nia serpentine endemic taxa present in a given region; assumption is incorrect at both taxonomic levels, ``restricted endemics'' includes only those taxa restricted but the root of this difference seems to be largely to a given region; black bars represent Kruckeberg's at the level of genus. A number of families con- (1984) estimates of restricted endemics. tribute a much higher proportion of the serpentine 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 229

TABLE 3. EXAMPLES OF ``REGIONAL'' SERPENTINE INDICATORS, SENSU KRUCKEBERG (1984). 1 CA ϭ California, KL ϭ Klamath Ranges, NC ϭ North Coast Ranges, BA ϭ San Francisco Bay Area, SC ϭ South Coast Ranges, SN ϭ Sierra Nevada.

Taxon Distribution1 Comments1 Allium amplectens CA SCÐbroad endemic; Northern CAÐweak indicator at best Aspidotis densa CA Marin CountyÐbroad endemic; rest of NCÐweak to strong indicator; KLÐbroad endemic to strong indicator, depending on locality; rest of CAÐstrong indicator Festuca californica CA Northern SN and KLÐstrong indicator to broad endemic; NCÐprimar- ily weak indicator; rest of CAÐindifferent Lupinus onustus KL, SN KLÐbroad endemic; SNÐindifferent (mostly non-ultrama®c) Pinus attenuata CA Mendocino County and neighboring NCÐbroad endemic; rest of NC and SCÐstrong indicator; KLÐweak indicator; SNÐweak indicator to indifferent Pinus jeffreyi KL, NC, SC, SN KL and NCÐϮ strict endemic; Westslope of northern SNÐstrong indi- cator; rest of CAÐindifferent Quercus vaccinifolia KL, NC, SN Mendocino County and neighboring NCÐbroad endemic; Northern NC and KLÐweak indicator; SNÐindifferent Sedum obtusatum KL, SN KL and NCÐϮ broad endemic; SNÐweak indicator or indifferent ssp. obtusatum Stachys pycnantha CA Marin CountyÐbroad endemic to strong indicator; Northern SNÐvery weak indicator; rest of CAÐweak indicator or indifferent Viola douglasii CA Plumas CountyÐendemic; NCÐstrong indicator; rest of CAÐindifferent endemic ¯ora than they do of the California endem- (2) lower mean serpentine af®nity scores, and (3) ic ¯ora (Fig. 1), but our database shows that most high standard deviations in their af®nity scores. Ta- of these ``anomalies'' are due to one or two genera ble 3 lists ten examples of regional indicator taxa within those families (see Fig. 2). Examples include in our database. Fritillaria and Allium in Liliaceae, Minuartia in Some of the variability in our serpentine af®nity Caryophyllaceae, Streptanthus and Arabis in Bras- scores is thus due to geographic variation in af®n- sicaceae, Hesperolinon in Linaceae, and Eriogon- ities, but some is also due to inadequate, statisti- um in Polygonaceae. Many of these genera are cally biased, or even faulty information from our well-known as foci of neoendemism (i.e., genera sources. We attempted to offset these sources of with groups of actively and rapidly speciating taxa) variability by including as many sources as possible (Raven and Axelrod 1978). It is interesting that in our database (and by using accession records), such prominent California plant families like Scro- but were not successful in all cases. We consider phulariaceae, Hydrophyllaceae, Boraginaceae, On- any taxon with a standard deviation in af®nity score agraceae and Polemoniaceae are underrepresented Ͼ 1.5, or having fewer than three sources, as being on serpentine substrates. Certain highly diverse in ``need of further research''; this includes about genera in California are also proportionally under- a third of the taxa in our database. Examples of represented as serpentine endemics (e.g., Clarkia, such taxa include: Lupinus lapidicolaÐcalled a Phacelia, Ceanothus, Gilia, and Mimulus). strict serpentine endemic by Kruckeberg (1984) and As we constructed our database, taxa with high a strong serpentine indicator by CNPS (2004), and variability in serpentine af®nity scores were tagged with 2/2 accession records in SMASCH with ultra- for further research (e.g., through accession rec- ma®c habitat descriptions, but stated as occurring ords; see Methodology) so that we might be able only on granites by the Jepson Manual (Hickman to discern taxa that truly varied geographically in 1993) and Munz and Keck (1973); Phacelia pha- their af®nities from taxa that simply suffered from celioidesÐKruckeberg (1984) and V. Yadon (per- inadequate or faulty information. The former were sonal communication) believe this is a strict endem- called ``regional indicators'' by Kruckeberg (1984), ic, but the Jepson Manual is mute on the subject, i.e., taxa that are considered serpentine endemics or and only 1/3 accession records in SMASCH are on indicators in one part of their range but show less ultrama®cs (but the two nonserpentine locations or no af®nity for ultrama®c substrates in other parts may have misidenti®ed geology given the location); of their range. In his Appendix D, Kruckeberg and Allium lacunosum var. lacunosumÐboth the (1984) tried to summarize where the different re- Jepson Manual and Kruckeberg rate this as a strict gional indicators he had identi®ed occurred on ul- endemic, Munz and Keck score it a strong indica- trama®cs. We refer the reader to Kruckeberg (1984) tor, but SMASCH has only 1/6 records on ultra- for details on these taxa (most of which also occur ma®cs. in our database), but most regional indicators in our Some species had surprising levels of ultrama®c database can be recognized by searching for taxa af®nity. For example, our database includes a num- with: (1) relatively wide geographic distributions, ber of taxa that we personally have only rarely seen 230 MADRONÄ O [Vol. 52 on serpentine (e.g., Lathyrus vestitus var. vestitus, should be a high priority for land management Apiastrum angustifolium, Emmenanthe penduli¯ora agencies and private conservation organizations var. penduli¯ora). It also includes other taxa which throughout the State. we would have characterized as being clearly in- Our database of serpentine af®nity updates, and different to ultrama®c substrates, but which scored expands on the widely-used tables of serpentine en- higher based on our sources (e.g., Adenostoma fas- demic and ``indicator'' taxa published in 1984 by ciculatum, Pinus balfouriana ssp. balfouriana). As Art Kruckeberg in his classic monograph on Cali- noted above, some of these ``discrepancies'' may fornia serpentine ecology. Our data are also a quan- be due to inadequate or biased dataÐthe ultrama®c titative synthesis of the qualitative (and usually in- af®nity of these types of taxa will drop as we col- complete) allusions to serpentine af®nity contained lect more information. Many of these surprising af- in habitat descriptions in California ¯oras and ¯ora ®nities are probably real however, and they are sim- databases, including Munz and Keck (1973), Hick- ply a sign of our limited knowledge of the relation- man (1993), Oswald (2002), the online CalFlora ships between California plant life and ultrama®c Database (CalFlora 2004), and the California Na- substrates. tive Plant Society Online Inventory of Rare and In accession records, and in the literature, bota- Endangered Plants (CNPS 2004). Our data on ser- nists and ecologists frequently misidenti®ed basic pentine endemism should prove valuable to efforts rock types. For example, in accession records we in ecology, biosystematics (Baldwin 1995), conser- found a number of examples of peridotite being vation, and land management. In particular, we called ``volcanics'' or even ``sandstones''. We also hope that our database will help us better under- found multiple examples, in accession records as stand the nature and degree of serpentine endemism well as in the literature, of gabbro and other basic in the California ¯ora, and we hope it will spur the intrusive rocks being misidenti®ed as ultrama®cs. collection of additional, critical information neces- Gabbro and ``basic'' rocks are ``ma®c'' in compo- sary for conserving the rare plants and habitats of sitionÐthat is to say, they usually contain visible ultrama®c substrates. feldspars and they are geochemically distinct from ultrama®c rocks. For example, the average alkali- ACKNOWLEDGEMENTS gabbro contains 4±5 times as much Na as perido- We would like to thank the following individuals for tite, 5±10 times as much P, 3±4 times as much K providing information on plant af®nities to ultrama®c sub- and Ca, and about ⅓ as much Mg (Ehlers and Blatt strates: Joe Callizo, Sydney Carothers, Susan Erwin, Lin- 1982). The famous gabbro outcrops of Eldorado nea Hanson, Lisa Hoover, David Isle, Marla Knight, Niall County (Pine Hill) or San Diego County are there- McCarten, John McRae, Julie Kierstad Nelson, Tom Nel- fore not ultrama®c, even though the effect of the son, Robert Preston, Barbara Williams, and Vern Yadon. substrate on plant physiognomy and community Dick Moe of the Jepson Herbarium provided us with data composition may appear similar. A number of spe- on identities and numbers of taxa endemic to California. cies in our database appear to be primarily, if not Special thanks to Art Kruckeberg and Niall McCarten for exclusively gabbro endemics, but we lacked suf®- comments on early versions of the manuscript. cient information to remove them from our list. These include Acanthomintha ilicifolia, Fremonto- LITERATURE CITED dendron californicum ssp. decumbens, and Calo- BALDWIN, B. G. 1995. A new prospect for California bot- chortus weedii var. vestus. any: integrating biosystematics and phylogenetics. As has been frequently noted (Mason 1946a, b; MadronÄo 42:154±167. Raven and Axelrod 1978; Kruckeberg 1984, 2002; . 1999. New combinations and new genera in the Skinner and Pavlik 1994; McCarten 1997), Cali- North American tarweeds. Novon 9:462±471 fornia's ultrama®c soils support a very high pro- . 2001. Harmonia guggolziorum (Compositae-Ma- portion of the State's rare plants. Based on our da- diinae), a new tarweed from ultrama®cs of southern tabase, almost 11% (111/1021) of California's rare Mendocino County, California. MadronÄo 48:293± 297. plant taxa are either broadly or strictly restricted to BARKLEY, T. M. 1999. The segregates of Senecio, s.l. and ultrama®c substrates; 15% of List 1b taxa (154/ Cacalia, s.l., in the Flora of North America North of 1021) show high af®nity for ultrama®c substrates Mexico. Sida 18:661±672. (i.e., they are endemics or indicators). In north- BROOKS, R. R. 1987. Serpentine and its vegetation. Dios- western California, 15% of plant taxa managed as corides Press, Portland, OR. ``sensitive'' by the Forest Service are serpentine en- CALFLORA. 2004. CalFlora Species Database (http:// demics, and fully 30% are closely associated with www.cal¯ora.org). CalFlora, Berkeley, CA. ultrama®cs (J. K. Nelson and L. Hoover personal CLIFTON, G. 2001. Plumas County and Plumas National communication). In 2002, Kruckeberg wrote that Forest ¯ora. Unpublished ¯ora printed by the Plumas National Forest, Quincy, CA. ``preservation of serpentine habitats in California is CNPS. 2004. California Native Plant Society Online In- spotty, inadequate, and largely coincidental''. Giv- ventory of Rare and Endangered Plants (http://www. en the great importance of ultrama®c substrates to cal.net/ϳlevine/cgi-bin/cnps/sensinv.cgi). 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es.html). University and Jepson Herbaria, University U.S. FISH AND WILDLIFE SERVICE. 1998. Draft recovery of California, Berkeley, CA. plan for serpentine soil species of the San Francisco UC-JEPS. 2004b. Jepson Flora Project (http://ucjeps. Bay Area. U.S. Fish and Wildlife Service, Portland, ࿞ ࿞ berkeley.edu/interchange/I index newtax.html). Uni- OR. versity and Jepson Herbaria, University of California, WAGNER, D. W. 1979. Systematics of Polystichum. Pter- Berkeley, CA. UC-JEPS. 2004c. Species Management System for Cali- ologia 1:1±64. fornia Herbaria (http://www.mip.berkeley.edu/www࿞ ZIKA, P. F., K. KUYKENDALL, AND B. WILSON. 1998. Carex apps/smasch/smasch࿞accession.html). University and serpentinicola (Cyperaceae), a new species from the Jepson Herbaria, University of California, Berkeley, Klamath Mountains of Oregon and California. Ma- CA. dronÄo 45:261±270. 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 233 ϭ 11 strong ϭ Lifeform parasitic, rhiz ϭ San Francisco Bay ϭ 10 Tax. Cat. Dicot Annual forb Dicot Perennial forb DicotDicot Perennial forb Dicot Perennial forb Dicot Perennial forb Perennial forb Sum of all af®nity scores, including 9 4 hemiparasitic, paras ϭ broad endemic/strong indicator, SI North Coast Ranges, BA ϭ 1111 Geog. Dist. ϭ Names as in Hickman (1993); names in parentheses 1 11111 11111 1111 11111 1111 KL NC BA SC SN 8 California Native Plant Society rarity codes, from CNPS 8 cespitose, hemipar Rarity ϭ 7 SE broad endemic, BE/SI 6 Klamath Ranges, NC ϭ SD ϭ 5 carnivorous, cesp . Ordered by family and taxon. ϭ carn Sources Med. 11 Standard error of af®nity scores. ALIFORNIA 7 strict endemic, BE C 4 ϭ Mean af®nity score, including information from CalFlora. Sum 3 3 Geographic distribution: KL 9 UBSTRATES IN Mean S Af®nity: SE 2 2 Taxonomic category. 10 LTRAMAFIC U weak indicator/indifferent. ϭ Sierra Nevada. Family Aff Standard deviation of af®nity scores. FFINITY TO 6 ϭ A ApiaceaeApiaceaeApiaceaeApiaceae WI/IN SEApiaceae 1.4 SEApiaceae BE/SIApiaceae 5.75 6.0Apiaceae 3.6 6.2 SEApiaceae 18 4 SEApiaceae 21.5 18.5 SEApiaceae 5.8 1.4 SIApiaceae 3 5.9 BE 6 3Apiaceae 1.3 34.5 6.1 WI/INApiaceae 29.5 0.7 SI 6.0 3.5 2.7 6.0 24.5 1.4 5.0 SI 6Apiaceae 0.0 1.9 0.0 SE 5Apiaceae 8 25 2.75 0.0 SI 0.8 4 3.3 0.0Apiaceae 6.0 1 3.2 6.0Apiaceae 1b 0.8 13 2 6.1 WI 6.0Apiaceae 3 1 5 0.4 12.6 BE 0.3 2.8Apiaceae 42.5 0.0 1.4 WI 0.2Apiaceae 4 3.0 11 1.7 6.0 SE 4 0.0Apiaceae 0.9 4 4 4.6 WI 1 7 0.6 1Apiaceae 1.4 4 2.1 0.6 8.5 1 BE/SI 3.0Apiaceae 23 3.0 1 0.3 1 0.6 5.6 1 WI 4 6.0Apiaceae 10.6 1 3.7 2.3 1.7 1 WI 2.4 1 1Apiaceae Dicot 5 22.5 0.0 1 WI 1.1 2.0Apiaceae 18.5 1.2 5 1.8 5 BE 5 0.0Apiaceae 1 1.0 4 2.4 1.8 4 WI 1 4Apiaceae Dicot Perennial Dicot 4 Dicot 6.0 2.3 forb 7.1 5 WI/IN 1.4 2.0 1.2Apiaceae 1 5.3 3.5 BE/SI 3 6.0 2.4 0.6 1.3 2.0 2.1 4.5 1 1 WI/IN 3.0 26.5 4 Perennial Perennial Dicot 3.5 1.0 1.1 Perennial forb forb WI 0.9 1 forb 1.0 1 1 2 2.3 1.3 3.75 8.5 SI 0.5 Dicot 4 10.5 1 2 2.0 1.2 1.0 5 Dicot 1.5 4 4 3 Perennial 2.3 1 1 1.5 forb 0.7 4 4 2.0 1 Dicot 3 2.1 6.0 2.6 Perennial Dicot 0.7 forb 1.0 7 1 0.0 Perennial 1.5 forb 1.0 1.3 12.75 3 3.0 0.7 0.0 Perennial 4 0.6 2.6 Dicot forb Perennial Dicot 0.6 forb Dicot 0.4 5 1b 1.0 3 4 1.3 0.3 1.5 2.0 Dicot Perennial 1.0 4 Perennial forb Perennial 1 forb 0.9 forb 1 1 2.2 3.2 Dicot 1.0 1.9 Perennial forb 1 1 Dicot 1 1 Dicot 1 1 Perennial forb Perennial 1 1 forb Perennial forb Dicot Dicot 1 1 1 Dicot Dicot 1 Perennial Perennial forb Dicot forb Dicot 1 1 Perennial Perennial forb Dicot forb 1 Perennial Perennial forb forb Dicot Perennial Dicot forb Perennial forb Perennial forb ApiaceaeApiaceae SI WI 2.7 1.5 8 7.6 5 3 0.1 3.0 2.5 1.5 1.1 0.9 1 1 1 Dicot Perennial forb IGH H ssp. var. AXA WITH T weak indicator, WI/IN 1 LANT ϭ South Coast Ranges, SN Median of af®nity scores. Taxon 5 ϭ 1. P triternatum dasycarpum PPENDIX CalFlora. are based on more recent revisions (see text for sources). indicator, WI Area, SC A rhizomatous. See text for more information. Inventory of Rare and Endangered Plants of California, 11-2004. Ligusticum californicum Lomatium ciliolatum Lomatium congdonii Lomatium dasycarpum Lomatium engelmannii Lomatium hooveri Lomatium howellii Lomatium macrocarpum Lomatium marginatum Lomatium observatorium Lomatium parvifolium Lomatium repostum Lomatium tracyi Lomatium triternatum Lomatium utriculatum Perideridia bacigalupii Perideridia kelloggii Perideridia leptocarpa Perideridia oregana Perideridia pringlei Sanicula bipinnati®da Sanicula hoffmannii Sanicula maritima Sanicula peckiana Sanicula tracyi Sanicula tuberosa Tauschia glauca Tauschia hartwegii Tauschia howellii Tauschia kelloggii Angelica tomentosa Apiastrum angustifolium 234 MADRONÄ O [Vol. 52 11 (rhiz.) (rhiz.) Lifeform 10 Tax. Cat. DicotDicot Annual forb Annual forb Dicot Annual forb Dicot Annual forb Dicot Annual forb Dicot Perennial forb 9 1111 1111 Geog. Dist. 11111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff AsclepiadaceaeAsteraceae SE WI/IN 6.0 1.4 42 4.1 7 3 6.0 1.0 0.0 1.5 0.0 0.9 4 1 Dicot Perennial forb AsteraceaeAsteraceaeAsteraceaeAsteraceae SI WIAsteraceae SE SE 3.3Asteraceae 1.9Asteraceae 13 SE 5.6 7.75 6.1 22.5Asteraceae WI/IN 24.5Asteraceae SI 5.5 4 4 1.1Asteraceae 4 16.5Asteraceae SE 4 0.8 3.0 3.25 BE/SI 2.5 6.0 2.7 WI/IN 0.5 3 6.0 3.7 15 3 SI 6.2 1.0 1.4 0.3 1.3 0.0 11 6.0 0.5 18.5 1.0 0.0 4 4 1.2 3.1 6 0.9 4 3 0.7 1 3 15.5 0.5 1 2.0 4 3 1 1 6.0 1 4.0 1.1 5 0.0 2.0 1 0.5 0.6 1 0.0 3.0 1.2 1b 0.3 1 1b 0.7 1.2 0.6 1 Dicot 1 Dicot 1 Dicot 1 1 1 Perennial 1 forb Perennial forb Dicot Dicot Perennial forb 1 Dicot Perennial Perennial forb 1 forb Dicot Dicot Perennial forb Dicot Perennial forb Perennial forb Perennial forb AsteraceaeAsteraceaeAsteraceae SI BE WI 2.6 5.3 18 2.1 21 12.5 7 4 6 2.0 5.5 2.5 1.6 1.0 1.1 0.6 0.5 0.5 1b 1 1 Dicot Dicot Annual forb Annual forb AsteraceaeAsteraceae WI SI 1.7 5.1 2.5 10 3 2.0 4 1.5 2.5 0.9 0.6 0.3 1 1 1 Dicot Perennial forb AsteraceaeAsteraceaeAsteraceae SEAsteraceae WI WI 6.1 1.8 SI 30.5 1.7 8.85 3.0 5 5 5 12 6.0 2.0 0.0 3 1.5 4 0.0 0.7 2.0 1b 2.0 0.9 1 2.0 0.5 1.0 1 1b 1 1 1 Dicot Dicot Perennial forb Shrub Dicot Perennial forb Asteraceae SI 3.0 12 4 3.0 1.6 0.8 1 1 1 Dicot Perennial forb Asteraceae SE 5.9 29.5 5 6.0 0.4 0.2 1b 1 1 Dicot Perennial forb AsteraceaeAsteraceaeAsteraceae SEAsteraceaeAsteraceae WI/INAsteraceae SI 6.1 1.0Asteraceae WI 24.5 WI/IN 3.1 WI 2.7 1.3 2.0 SE 4 8 3 10 4 2.0 6.0 5.5 1.0 8.1 0.0 38.5 3 5 1.0 3 0.0 0.5 4 3.0 1b 2.0 1.0 7 0.6 0.7 1.5 1.5 6.0 0.3 0.3 0.9 2.1 1 1 1.0 1.1 1 1 0.4 1 4 1 1 1 1 1 1 1 1 Dicot 1 Dicot 1 1 Perennial 1 1 forb 1 Perennial Dicot Dicot forb Dicot 1 Dicot Dicot Shrub Annual Perennial forb forb Shrub Shrub Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Perennial forb Asteraceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 1 Dicot Perennial forb ssp. var. var. var. . breweri campylon fontinale var. var. var. var. var. 1 ONTINUED Taxon 1. C macrolepis glabriuscula heterocarpha vaseyi obispoense PPENDIX Asclepias solanoana Agoseris heterophylla Ancistrocarphus ®lagineus Antennaria argentea Antennaria suffrutescens Arnica cernua Arnica spathulata Aster oregonensis Balsamorhiza macrolepis Balsamorhiza sericea Brickellia greenei Cacaliopsis nardosmia Calycadenia multiglandulosa Calycadenia oppositifolia Calycadenia pauci¯ora Calycadenia truncata Chaenactis glabriuscula Chaenactis glabriuscula Chaenactis suffrutescens Chrysothamnus nauseosus Cirsium andrewsii Cirsium cymosum consimilis Cirsium fontinale Cirsium douglasii Cirsium fontinale Cirsium remotifolium Coreopsis stillmanii Crepis pleurocarpa Ericameria arborescens Ericameria greenei Ericameria ophitidis Cirsium hydrophilum Cirsium fontinale A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 235 11 (rhiz.) (rhiz.) (rhiz.) forb, Shrub succulent) forb, Shrub forb, Shrub Perennial forb Lifeform 10 Tax. Cat. Dicot Shrub 9 Geog. Dist. 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff AsteraceaeAsteraceae WI BE 1.7 4.8 5 28.5 3 6 2.0 6.0 1.5Asteraceae 2.1 0.9Asteraceae 0.8Asteraceae 4 WI WI/INAsteraceae 1.2 1.8 WI 3.6 WI 5.25 1.7 3 3 1.8 5 2.0 1.0 1 5.25 1 1.4 1.0 Dicot 3 Dicot 3 0.8 0.5 2.0 2.0 Perennial forb Perennial 0.9 forb 1.4 0.5 0.8 1b 1 1 1 1 1 1 1 Dicot Dicot 1 Perennial Perennial forb forb Dicot Dicot Perennial forb Perennial Asteraceae BE/SI 3.7 11 3Asteraceae 3.0 1.2Asteraceae 0.7 WIAsteraceae WI 2.3 SE 1 1.7 7 5.5 5 3 16.5 3 2.0 3 0.6 2.0 6.0 0.3 Dicot 1.5 1.2 0.9 0.7 Perennial forb 1b 1 1 1 1 Dicot Dicot Shrub Shrub AsteraceaeAsteraceaeAsteraceae SEAsteraceae SE SI 5.7 WI 6.2 28.5 3.3 18.5 1.5 10 5Asteraceae 3 4.5 6.0Asteraceae 3 6.0Asteraceae WI 0.9 3Asteraceae 0.0 4.0 0.4 WI 0.0Asteraceae 2.0 2.4 1b SE 3.1 2 WI 1.2 1.8 2.0 9.5 0.7 BE/SI 6.2 1 1 1.9 8 4 18.5 4 3.5 1 3.75 17.5 2.0 3 4 2 0.5 1 5 6.0 2.0 0.3 1.9 0.0 1.6 3.0 Dicot 4 1.6 0.0 0.8 1.5Asteraceae Dicot 1.1 1Asteraceae 1b 0.7 Perennial forb Dicot 4Asteraceae Perennial SE Dicot forb 1 WI 1 1 Perennial forb 6.0 WI Perennial forb 2.0 18 1.9 6.1 1 Dicot 1 1 5.75 3 3 Dicot Dicot 1 6.0 3 Perennial forb 3.0 Dicot Dicot 0.0 2.0 1.7 Perennial Perennial forb forb 0.0 1.0 1.1 Shrub Shrub 1b 0.7 1 1 1 Dicot Dicot Dicot Annual forb Shrub (stem Perennial AsteraceaeAsteraceae SI SE 2.9 5.7 11.5 45.5 4 8 2.5 6.0 2.3 1.1 1.2 0.4 4 1 1 1 1 1 1 1 1 Dicot Dicot Perennial Annual, forb Asteraceae WI/IN 1.0 2 2 1.0 1.4 1.0 1 Dicot Perennial var. . davyi nudatus whitneyi con®nis var. var. var. var. var. var. var. var. var. var. var. var. var. 1 ONTINUED Taxon 1. C sierrensis hirsutula maritima de®ciens achillaeoides lanceolatum robustior discoidea viscidulus tanaceti¯orum PPENDIX Erigeron petrophilus Grindelia hirsutula Grindelia hirsutula Gutierrezia californica Erigeron lassenianus Eriophyllum latilobum Grindelia hirsutula A Erigeron angustatus Erigeron bloomeri Erigeron cervinus Erigeron decumbens Erigeron foliosus Erigeron petrophilus Erigeron reductus Erigeron serpentinus Eriophyllum conferti¯orum Eriophyllum jepsonii Eriophyllum lanatum Eriophyllum lanatum Harmonia guggolziorum Hazardia stenolepis Hazardia whitneyi Hazardia whitneyi Helenium bigelovii Helianthus exilis 236 MADRONÄ O [Vol. 52 11 forb, Shrub Lifeform 10 Tax. Cat. Dicot Perennial Dicot Annual forb Dicot Annual forb DicotDicot Annual forb Annual forb Dicot Annual forb 9 1111 Geog. Dist. 11111 11111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff AsteraceaeAsteraceaeAsteraceae WI/IN 1.3Asteraceae SI BE 4Asteraceae BE 2.5 5.1 WI 3 7.5 30.5 5.3 1.0 31.5 2.0 3 6 1.5 6 3.0 6 0.9 5.5 1.2 1.3 5.5 0.7 0.5 3 1.0 3 1b 0.4 2.0 1b 1.0 0.6 1 1 1 1 Dicot Dicot Dicot Annual Annual forb forb Annual forb AsteraceaeAsteraceaeAsteraceae WIAsteraceaeAsteraceae BE/SIAsteraceae 2.0 WI 3.8Asteraceae WIAsteraceae BE 6 15Asteraceae 2.2 SE 2.0 BE/SI 4.7 SI 6.5 3 3.5 4 SE 6.1 6.1 23.5 10.5 3.0 24.5 3 4.5 3.2 3 6.2 1.7 5 2.6 3.0 19 3 4 1.0 18.5 3.0 1.3 5.0 1.4 3.0 1.7 6.0 0.8 1.7 6 3 0.6 1.0 0.0 0.7 1 0.3 1 3.5 0.0 6.0 1 1b 1 1b 1.4 0.0 1 0.6 0.0 1 1 1b 1b 1 1 Dicot 1 Dicot 1 1 1 1 1 Dicot Perennial forb Dicot Perennial Dicot forb Dicot Dicot Perennial forb Annual forb Annual forb Annual Dicot Dicot Annual forb forb Annual Annual forb forb AsteraceaeAsteraceae WIAsteraceaeAsteraceae WI/INAsteraceae 1.5 1.3 WI SI 4.5 4 WI 1.8 3 3.0 1.8 5.25 3 2.0 12 7.25 3 2.0 1.2 1.2 0.7 4 4 2.0 0.7 4 0.7 1.5 3.0 0.4 1.6 2.4 4 0.8 1.2 1 1 1 1 1 1Asteraceae Dicot 1 Dicot 1Asteraceae 1 BE Annual forb Dicot Annual forb WI Dicot 5.4 Dicot Annual forb 32.5 1.8 Annual forb Annual forb 7.25 6 4 5.5 0.8 2.0 0.3 1.4 1b 0.7 1 Dicot Annual forb AsteraceaeAsteraceaeAsteraceae BE/SI BE 4.1 WI/IN 16.5 1.4 5.4 27 4.25 4 3 4.0 5 1.8 2.0 6.0 0.9 1.0 1.3 4 0.6 0.6 1 1 1 1 1 Dicot 1 Dicot Dicot Annual forb Perennial forb Annual forb AsteraceaeAsteraceae SE SI 6.1 42.5 3.0 7 9.1 6.0 3 0.0 3.0 0.0 3.0 1b 1.7 1 Dicot Annual forb AsteraceaeAsteraceaeAsteraceae WI WI WI/IN 2.1 1.3 2.4 6.25 4 7.25 3 3 3 3.0 1.0 1.6 1.0 3.1 0.9 0.6 1.8 0.3 1 1 1 Dicot Annual forb AsteraceaeAsteraceae WI WI 2.4 2.4 4.75 7.25 2 3 2.4 3.0 2.3 1.3 1.6 0.8 1b 1 1 1 Dicot Dicot Annual forb Annual forb ) Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Annual forb tracyi . var. var. ϭ var. var. var. ( ssp. ssp. ssp. ) 1 Harmonia h. ONTINUED ) ϭ ( Taxon 1. C Harmonia s. ϭ ( glabrata micradenia californica oregona Harmonia d. calyculata congesta sparsi¯ora PPENDIX Madia stebbinsii Lessingia hololeuca Lessingia micradenia Lessingia micradenia Lessingia nemaclada Hieracium bolanderi Hieracium greenei Lagophylla glandulosa Lagophylla minor Layia discoidea Layia jonesii Layia septentrionalis Lessingia arachnoidea Lessingia ®laginifolia Hemizonia congesta Hemizonia congesta Hemizonia congesta Hemizonia halliana Hesperevax sparsi¯ora Heterotheca oregona Madia exigua Lessingia occidentalis Lessingia ramulosa Luina hypoleuca Madia doris-nilesiae Madia hallii Malacothrix clevelandii Malacothrix ¯occifera Micropus amphibolus Microseris douglasii Monolopia gracilens Pentachaeta bellidi¯ora A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 237 11 (rhiz.) (rhiz.) Lifeform 10 Tax. Cat. Dicot Annual forb Dicot Perennial forb 9 Geog. Dist. 11111 1111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff BoraginaceaeBoraginaceae BE/SI 4.4 WI 17.5 1.5 4 3 4.5 2.1 2 1.0 1.5 1b 2.1 1.5 1 1 Dicot Annual forb Dicot Annual forb Asteraceae BE/SI 3.8 15 4 3.0 1.5 0.8 1 1 Dicot Perennial forb AsteraceaeAsteraceae SEAsteraceae BE/SIAsteraceae 6.2 4.0Asteraceae WI 18.5 16 SE 1.7 WI 3 4 6.0 5 6.0 1.9 30 4.5 0.0 7.5 3 0.0 2.4 1.2 2 5 1.0 4 2.1 6.0 1 2.0 1.2 0.0 1 1.3 0.0 0.7 1b 1 1 1 Dicot 1 Dicot Perennial forb Dicot Perennial forb Dicot Perennial forb Perennial forb Asteraceae BE 5.3 21 4 6.0 1.5 0.8 1 1 Dicot Perennial forb Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Perennial forb Asteraceae SE 5.8 40.5 7 6.0 0.8 0.3 1b 1 Dicot Perennial forb Berberidaceae WI 1.7 5 3 1.0 1.2 0.7 BerberidaceaeBerberidaceae WI SE 1.7 6.2 5 18.5 3 3 6.0 1.0 0.0 1.2 0.0 0.7 4 1 1 1 1 1 Dicot Dicot Perennial Shrub forb AsteraceaeAsteraceaeAsteraceae BEAsteraceaeAsteraceae SEBerberidaceae 5.1 WI/IN WI 20.5 1.1 WI 6.2 WI 18.5 2.1 1.8 4 1.6 1.5 5.5 3 6.0 2 4.75 3 2.0 6.0 1.1 3 3 1.0 0.0 1.3 2 2.0 4 1.0 0.0 1.0 0.6 1.5 1.2 4 1 0.3 0.7 0.7 1b 0.5 1 1 1 1 1 Dicot 1 1 Dicot Dicot 1 Perennial forb Dicot Dicot 1 Perennial Perennial forb forb Annual Dicot forb Shrub Perennial forb Asteraceae SE 5.8 46.5 8 6.0 0.7 0.3 4 1 Dicot Perennial forb ) Asteraceae BE 5.3 32 6 6.0 1.6 0.7 1 1 Dicot Perennial forb ) Asteraceae BE 4.9 29.5 6 5.0 1.3 0.5 1b 1 Dicot Perennial forb ) repens . var. var. var. var. var. var. var. var. var. Packera l. Packera g. Packera c. var. var. Packera lewisrosei ϭ Packera ϭ ϭ ) ϭ 1 ( Packera ( ( ( ONTINUED var. var. ϭ ) ( ϭ ( Taxon 1. C ) ) Packera m. ϭ dissita ( eurycephala eurycephala lewisrosei congesta pinetorum racemosa glauca clevelandii v. h. eurycephalus eurycephala aquifolium c. v. c. heterophyllus PPENDIX Cryptantha clevelandii Cryptantha excavata Senecio macounii Senecio eurycephalus A Pyrrocoma racemosa Pyrrocoma racemosa Pyrrocoma racemosa Raillardella pringlei Rigiopappus leptocladus Senecio clevelandii Rudbeckia californica Senecio eurycephalus Senecio greenei Senecio layneae Vancouveria planipetala Vancouveria chrysantha Solidago guiradonis Solidago multiradiata Stebbinsoseris decipiens Wyethia bolanderi Berberis aquifolium Berberis aquifolium Senecio clevelandii 238 MADRONÄ O [Vol. 52 11 (rhiz.) (rhiz.) (rhiz.) (rhiz.) Lifeform 10 Tax. Cat. DicotDicot Annual forb Annual forb Dicot Annual forb 9 Geog. Dist. 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff BrassicaceaeBrassicaceae BEBrassicaceae BE 5.2Brassicaceae WI 15.5 5.4Brassicaceae SE 48.5Brassicaceae 2.0 3Brassicaceae SIBrassicaceae 6 6.2 SE 9 5.0Brassicaceae WI/IN 18.5Brassicaceae 1.0 SI 6.0 2.7 1.4 6.1 WI 0.6 3Brassicaceae 1.0 BE 3 8 24.5 1b 4.25 0.3 3.0 2.0Brassicaceae 2.3 BE/SI 6.0 1 3Brassicaceae 3 1.0 5.3 4 9 3Brassicaceae 0.0 4.3 9.25 SE 0.6 31.5 1.0 SE 0.0 6.0 34.5 3.0 SE 4 1.4 3 1b 0.0 1 5.6 6 0.6 0.8 6.0 8 2.5 0.0 28 0.3 3.0 6.1 6.0 24 1b 1.6 1b 4.5 0.0 1 24.5 1.3 0.8 1 1.9 0.0 Dicot 5 1 1 0.5 1 4 0.7 4 4 1b Dicot 6.0 1 1b Perennial 6.0 forb 6.0 0.5 1 0.0 Perennial 0.0 0.2 forb Dicot 1 1 Dicot 0.0 0.0 1 1 1 4 1b 1 1 1 Perennial forb Dicot 1 Annual 1 forb Dicot Dicot 1 1 1 Dicot Dicot Perennial forb Dicot Perennial Perennial forb forb 1 Dicot Annual Perennial forb forb Annual forb Annual forb Dicot Dicot Dicot Annual Perennial forb forb Annual forb Boraginaceae WI 1.6 4.75 3 2.0 0.7Brassicaceae 0.4 Brassicaceae BE/SIBrassicaceae 3.9 SI 27 WI 2.9 7 17.5 1.9 3.0 3.75 6 1.6 2 2.5 0.6 1.9 1.9 0.8 1.6 1.1 1 1 1 Dicot 1 1 Dicot Perennial forb Dicot Perennial forb Perennial forb BrassicaceaeBrassicaceaeBrassicaceae SEBrassicaceae BE BE/SI 5.5 SI 3.8 5.4 16.5 11.5 32.5 3.2 3 16 3 6 6.0 3.0 6.0 1.2 5 2.1 1.0 0.7 1.2 0.4 3.0 1b 4 1b 1.6 1 0.7 1 1 1 1 1 1 Dicot Dicot Dicot Perennial forb Perennial forb Perennial forb Dicot Perennial forb BoraginaceaeBoraginaceaeBoraginaceae SEBoraginaceae WI/INBoraginaceae SE 1.4Boraginaceae 6.0 SIBoraginaceae WI/IN 24 4.1Brassicaceae 6.2 WI/IN 1.4Brassicaceae WI/IN 3.3 18.5 1.3 3 SE 1.3 4.1 4 9.75 SE 4 3 1.0 2.5 6.0 6.1 3 3 1.5 6.0 5.9 0.0 24.5 3 0.9 2 1.0 3.0 0.0 0.0 41.5 1.5 2.6 1.0 0.0 1.3 4 0.9 1.5 7 1.5 1b 1.1 6.0 0.9 1 0.8 6.0 0.0 1 0.4 0.0 1 1 0.1 2 1 1 1b 1 1 1 Dicot 1 Dicot 1 Dicot Dicot Annual forb Annual Dicot forb Annual forb Perennial 1 forb Dicot Dicot Perennial forb Perennial forb Perennial forb var. var. var. . var. ssp. ssp. var. var. stipitata var. 1 var. ONTINUED Taxon 1. C dissectifolia pachystigma barbarae gemmata albidus peramoenus horizontalis suffrutescens cuneata PPENDIX Cardamine nuttallii Cardamine pachystigma Cardamine pachystigma Caulanthus amplexicaulis Draba aureola Draba carnosula Draba howellii Erysimum franciscanum Guillenia ¯avescens Streptanthus albidus Streptanthus albidus Streptanthus barbatus Streptanthus barbiger Streptanthus batrachopus Cryptantha ¯accida Arabis suffrutescens Cardamine californica Arabis macdonaldiana Arabis oregana Arabis subpinnati®da Arabis suffrutescens Cryptantha hispidula Cryptantha intermedia Cryptantha mariposae Cryptantha milobakeri Cryptantha sobolifera Hackelia bella Pectocarya pusilla Arabis aculeolata Arabis constancei Arabis koehleri A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 239 11 Perennial forb Perennial forb Perennial forb Perennial forb Perennial forb Perennial forb Perennial forb Lifeform 10 Tax. Cat. Dicot Annual forb 9 Geog. Dist. 1111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Brassicaceae SEBrassicaceae SE 5.6Brassicaceae 22.5 SE 6.1 4 24.5 5.7 6.0 4 40 1.0 6.0 0.5 1b 7 0.0 0.0 6.0 1b 0.8 1 0.3 1 Dicot Dicot Annual, Annual, BrassicaceaeBrassicaceae SEBrassicaceae SEBrassicaceae WI 6.1Brassicaceae 24.5 BE 6.1 1.9Brassicaceae 36.5 SI 4 4.9 5.75Brassicaceae SI 6Brassicaceae 6.0 24.5 3 3.3 SE 0.0Brassicaceae 6.0 20 BE/SI 5 3.0 2.0Brassicaceae 0.0 0.0 4.0 SI 1.1 6.1 1b 3 6.0 0.0 SE 6 20 0.7Brassicaceae 30.5 1.8 4 3.3 0.8 3.0 1 1 SE 6.1 5 1Brassicaceae 5 16.5 1b 1.5 30.5 1 3.0 6.0 0.6 SE 4.0 6.1Brassicaceae 5 Ð 1 0.0 5 2.4 24.5 1 1 Ð 0.0 2.0 SE 6.1 1.1 1Brassicaceae 6.0 1 1b 1b 2.7 4 1 DicotBrassicaceae 24.5 0.0Brassicaceae 1.2 1 1 SE Dicot 6.1 6.0 0.0 SE 1b 4 Dicot 1 30.5 Annual 1 WI forb 1b 0.0 Dicot 6.1 Annual 6.0 forb 0.0 5.7 5 Annual 30.5 forb 1.6 1b 0.0 28.5 1 Annual forb 6.0 0.0 Dicot 8.2 5 1 1b 0.0 5 1 1 Dicot Dicot 6.0 0.0 5 Annual forb 6.0 Dicot 1b 0.0 2.0 1 0.9 0.0 Annual Perennial Dicot forb forb 1.6 Dicot 0.4 1b Annual forb 0.7 1 Annual forb Dicot Annual, 1 1 Dicot Annual, 1 Dicot Annual, 1 Dicot Annual, 1 1 Dicot Dicot Annual forb Annual forb Annual, lyonii ssp. ssp. ssp. ssp. . var. var. ssp. ssp. ssp. ssp. var. var. var. ssp. ssp. 1 hoffmanii ONTINUED var. Taxon 1. C brachiatus hoffmanii breweri glandulosus pulchellus secundus secundus elatus hirti¯orus kruckebergii morrisonii hesperidus insignis suffrutescens PPENDIX A Streptanthus brachiatus Streptanthus brachiatus Streptanthus breweri Streptanthus breweri Streptanthus drepanoides Streptanthus glandulosus Streptanthus glandulosus Streptanthus glandulosus Streptanthus glandulosus Streptanthus howellii Streptanthus insignis Streptanthus insignis Streptanthus morrisonii Streptanthus morrisonii Streptanthus morrisonii Streptanthus morrisonii Streptanthus niger Streptanthus polygaloides Streptanthus tortuosus 240 MADRONÄ O [Vol. 52 11 Perennial forb Perennial forb (rhiz.) (rhiz.) Perennial forb Lifeform 10 Tax. Cat. DicotDicotDicot Perennial forb Annual forb Annual, Dicot Annual forb DicotDicot Perennial forb Annual forb 9 Geog. Dist. 11111 11111 11111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Caryophyllaceae WI/INCaryophyllaceae WI 1.4 4.1 2.1 8.5 3 2.0 4 1.1 0.9 0.6 2.6 1.3 1 Dicot Perennial forb Brassicaceae SIBrassicaceaeBrassicaceae SE 3.3Campanulaceae BE/SICampanulaceae BE/SI 10Campanulaceae 4.4 BE/SI 6.1Campanulaceae 3.9 SE 22Campanulaceae 30.5 3.9 BE 19.25 3 SI 19.5Campanulaceae 6.0 5 3.0 5Campanulaceae 5.0 5 SE 18 WI/IN 5 0.6 2.5 15 6.0 4.0 4.0 0.3 1.0Campanulaceae 10 6.2 0.0 4.0 2.4 1.5 WI 3 4 0.0 18.5Campanulaceae 5 1.1 1.5 3 0.7 BE/SI 1b 6.0 0.7 4 1Campanulaceae 1.6 6.0 3 1b 0.0 3.8 1 BE 5 2.5 1.7Campanulaceae 3.25 0.0 19 1 6.0 1 SE 1.3 1.0 0.0 5.3 1 0.0 1 2 0.6 1.7 0.0 21 1 5 4 6.0 0.8 1.6 1 1b 1 18 1.9 3.0 1 1 Dicot 1Caryophyllaceae 4 1.4 2.0 SECaryophyllaceae 1 Dicot SI 1 0.9 3 5.5 Dicot Dicot 1 Annual, Dicot 6.1 1.0 1 6.0 Perennial 3.0 forb 0.5 Annual 24.5 1 Perennial forb 0.0 Annual forb Dicot forb 15 4 0.0 Dicot 4 Dicot Dicot Annual forb 5 6.0 Perennial Dicot forb 0.0 Perennial Annual 3.0 forb forb 1 0.0 1 0.7 Dicot 1 Perennial forb 1b 1 0.3 Dicot 1 1 1 Annual forb 1 Dicot Annual forb Dicot Annual forb Annual forb Dicot Perennial forb Brassicaceae WI/IN 1.4 4.25 3 2.0 1.0 0.6Campanulaceae WI/IN 1 1.0 1 2 1 Dicot 2 1.0 Annual, 1.4 1.0 1 Dicot Annual forb Caryophyllaceae WI 1.8 3.5 2 1.8 1.8 1.3 Caryophyllaceae WI 1.7 5Caryophyllaceae SECaryophyllaceae SICaryophyllaceae 3 SE 5.7Caryophyllaceae SECaryophyllaceae 3.2 2.0 28.5 SICaryophyllaceae 16 6.1 WI/IN 1.5 6.1 5 0.9 30.5 1.1 2.7 30.5 5 6.0 3.25 5 8 0.9 5 3.0 3 6.0 0.4 1.9 6.0 3 1b 0.0 1.0 0.9 0.0 0.0 0.9 1 3.0 0.0 4 0.5 0.6 1b 1 0.3 1 1 1 1 Dicot Dicot Dicot Perennial forb Dicot Perennial forb Perennial forb Perennial forb . var. (new candida glabrescens ssp. ssp. ssp. var. ssp. ssp. 1 glabra var. ONTINUED var. Taxon 1. C montanum pulchella serpentinicola campestris tortuosus taxon) gregaria PPENDIX Cerastium arvense Thelypodium brachycarpum Thlaspi californicum Thlaspi montanum Campanula angusti¯ora Campanula exigua Campanula grif®nii Campanula rotundifolia Campanula scabrella Campanula sharsmithiae Campanula wilkinsiana Githopsis diffusa Githopsis pulchella Githopsis pulchella Nemacladus montanus Arenaria kingii Minuartia decumbens Minuartia douglasii Githopsis pulchella Streptanthus tortuosus Minuartia californica Minuartia cismontana Minuartia howellii Minuartia nuttallii Minuartia rosei Minuartia stolonifera Moehringia macrophylla Silene antirrhina A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 241 11 (rhiz.) (rhiz.) Lifeform 10 Tax. Cat. Dicot Annual forb Gymnosp. Tree 9 Geog. Dist. 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff CrassulaceaeCrassulaceae BE/SI SI 4.0 16 3.2 16 4 3.5 5 1.4 3.0 0.7 2.2 1.0 1 1 1 Dicot 1 Dicot Perennial forb Perennial forb Crassulaceae WI 2.0 6 3 2.0 2.0 1.2 Cupressaceae SI 3.0 9 3 3.0 0.0 0.0 Cupressaceae BE/SI 4.0 8 2 4.0 2.8 2.0 1 Gymnosp. Shrub CupressaceaeCupressaceaeCupressaceae SICupressaceae SI BE BE 2.6 3.0 13 4.7 15 4.9 28 34 5 5 6 3.0 7 3.0 0.5 4.5 0.7 5.0 0.2 1.2 0.3 1.2 4 0.5 0.5 1 1 1 1 1 1 1 1 Gymnosp. 1 Tree Gymnosp. Gymnosp. Gymnosp. Tree Tree Tree Crassulaceae BE/SI 3.5 10.5 3 3.0 0.6 0.3 4 1 1 Dicot Perennial forb Crassulaceae SI 3.1 18.5 6 3.0 0.6 0.3 4 1 1 Dicot Perennial forb Caryophyllaceae BECaryophyllaceae BE/SI 5.3Caryophyllaceae 3.8 WICaryophyllaceae 31.5 BECaryophyllaceae 19 SICaryophyllaceae 1.8 SE 6 4.5Cistaceae 5 5.5 5.5 3.0 18 6.0Convolvulaceae 1.0 3.0 12Convolvulaceae BE 3 WI/IN 6 0.4 SE 1.3 4Convolvulaceae 1.0 4 2.0 0.6 4.7 4 BE 4.5Convolvulaceae 1.3 5.6 1 2 33 1Convolvulaceae BE 2.5 1.7 0.7 33.5Convolvulaceae 4.5 WI 1 6.0 2.2 0.9 1Crassulaceae BE/SI 18 2 7 4.9 ÐCrassulaceae 1.1 6 1 3.7 1.5Crassulaceae SE 24.5 Ð 1.0 1 6.0 SE 6.0 14.75 4 1Crassulaceae 4.5 1.4 SI 1.6 1.2Crassulaceae 5 6.2 1 1 1.0 4 0.6 4.5Crassulaceae 6.2 SE 0.5 Dicot 3 1 18.5 1Crassulaceae 3 5.0 WI 1.7 3.2 4.0 4 18.5Crassulaceae WI Dicot 1.0 1.3 0.9 2.4 6.1 SE 3 9.5 Perennial forb 2.0 0.6 1.3 SE 3 1.2 24.5 1.7 6.0 4 0.8 Perennial Dicot forb 1 1 4 3 8.1 6.1 6.0 Dicot 0.0 5.1 6.1 4 1 42.5 Dicot 0.0 3.0 0.0 4 24.5 Perennial 1 forb 0.0 6.0 Perennial 0.0 3 1b forb Dicot 7 1.5 1b 0.0 1 Perennial 0.0 forb 4 2.0 1 2.1 0.0 6.0 Dicot 1b Dicot Perennial Dicot 1.5 6.0 forb 1 1 1 1.1 1b 0.0 0.9 0.0 1 0.0 Shrub Perennial forb Dicot Perennial 0.0 1 forb 1b Dicot Dicot 1 1b Dicot 1 Perennial 1 forb 1 Annual Perennial 1 forb forb Dicot 1 1 Perennial 1 forb 1 Dicot 1 Dicot Perennial forb Dicot Perennial forb 1 Dicot 1 Perennial forb Perennial Dicot forb Dicot Dicot Annual forb Annual Perennial forb forb Perennial forb (in jackii . collina venusta bettinae murina ssp. (new var. ssp. ssp. bolanderi hookeri ssp. ssp. ssp. ssp. laxum heckneri ¯avidum ssp. ssp. ssp. 1 in Jepson) ONTINUED ssp. ssp. ssp. ssp. ssp. Taxon 1. C obtusatum oxyphylla tridactylosa campanulata glandulosa taxon) H. scoparium blochmaniae PPENDIX Sedum obtusatum Sedum radiatum Calocedrus decurrens Cupressus bakeri Cupressus lawsoniana Cupressus macnabiana Cupressus sargentii Juniperus communis Sedum laxum Sedum laxum A Silene campanulata Silene campanulata Silene grayi Silene hookeri Silene hookeri Silene serpentinicola Helianthemum suffrutescens Calystegia collina Calystegia collina Calystegia collina Calystegia collina Calystegia malacophylla Convolvulus simulans Dudleya abramsii Dudleya abramsii Dudleya blochmaniae Dudleya setchellii Parvisedum pentandrum Parvisedum pumilum Sedum albomarginatum Sedum eastwoodiae Sedum laxum 242 MADRONÄ O [Vol. 52 11 gram. (cesp.) gram. (rhiz.) (rhiz.) (rhiz.) gram. (cesp.) gram. (rhiz.) gram. (rhiz.) gram. (cesp.) gram. (cesp.) gram. (rhiz.) Lifeform 10 Tax. Cat. Monocot Perennial 9 1111 Geog. Dist. KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Cyperaceae SIDryopteridaceae SEDryopteridaceae WI 2.8Ericaceae 6.0 8.25 24 1.7Ericaceae 3 SEEricaceae 5.1 4 2.0 SEEricaceae 5.5 3Ericaceae 2.9 6.0 SI 27.5 1.7 6.3 0.0 2.0Ericaceae BE 0.0 12.5 1.5 SE 2.5 5Ericaceae 0.9 12.5 4.5 BE 2 6.0 6.2 22.5 1 1.3 SE 6.0 5 18.5 4.9 1 0.6 1 0.0 5 19.5 3.0 1b 6.2 1 3 0.0 1 1.5 4.0 18.5 1b 4 6.0 0.7 1.1 1 1 0.0 1b 3 0.5 4.5 Monocot Pteridoph. 0.0 1 1 4 1.0 1 6.0 Perennial Perennial 1a forb Pteridoph. 0.5 1 0.0 1 Perennial forb 1b 0.0 1 1b Dicot 1 Dicot 1 Shrub 1 Dicot 1 Shrub 1 Dicot Dicot Shrub 1 Dicot Shrub Shrub Dicot Shrub Shrub Cyperaceae WI/INCyperaceae 1.4 BECyperaceae 4.25 4.5 3 BE/SICyperaceae 22.5 3.8 2.0 BECyperaceae 23 1.0 5 0.6 4.9 SECyperaceae 4.0 6 24.5 1.7 3.5 BE 5.5 0.7 1 5 1.2 11 4 1 4.9 0.5 6.0 1 39 1.6 2 1 0.7 1 5.5 1 1b 8 Monocot 0.7 1 5.0 0.5 Perennial 1.1 2 1 0.4 Monocot 1 1 Perennial 1 Monocot Perennial Monocot Perennial Monocot Perennial Cyperaceae SI 2.6 10.5 4 2.3 2.9 1.4 1 Monocot Perennial ssp. . ssp. ssp. ssp. ssp. ssp. (new 1 ONTINUED Taxon 1. C bakeri sublaevis sonomensis franciscana montana ravenii taxon) PPENDIX Carex spissa Polystichum lemmonii Polystichum scopulinum Arctostaphylos bakeri Arctostaphylos bakeri Arctostaphylos canescens Arctostaphylos hispidula Arctostaphylos hookeri Arctostaphylos hookeri Arctostaphylos hookeri Carex brainerdii Carex gigas Carex mendocinensis Carex obispoensis Carex serpentinicola Carex serratodens Carex amplectens A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 243 11 (rhiz.) Lifeform 10 Tax. Cat. Dicot Annual forb Dicot Annual forb 9 Geog. Dist. 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff EricaceaeEricaceaeEricaceaeEricaceae BE/SI SIEricaceae 3.9 SE SI 19.5Ericaceae 2.8 5.7 BEEricaceae 5 2.6 5.5 28.5 WIEricaceae 10.5 5.0 4.0Fabaceae 2 WI 5 25 1.8 2.2Fabaceae 4 BE/SI 2.5 0.8 6.0Fabaceae 10.75 2.0 SIFabaceae 1b 0.7 3.0 3.5 5 0.9Fabaceae 5 6 0.5 1.7 SE 0.4 1 3.5 5.0 WI 3.2 4Fabaceae 0.9 3.0 4 WI/IN 1.0 15.75 1b 6.1 BE/SI 1.2 3 1Fabaceae 1 1.3 0.4 1.8Fabaceae 0.5 24.5 4.3 5 BE 3.0Fabaceae 3.0 3.75 3.5 25.5 1 1.7 Ð SI 3.0 4 3 4.6 1 SE 1.0 2 2.0 Ð 1 6 BE 6.0 1 Dicot 23 1 1.0 0.9 3 2.5 1.8 0.0 6.1 4.0 4 0.7 5.3 1.8 0.0 5 1 1 Shrub 24.5 1.2 5 Dicot 0.4 Dicot 10.5 1.3 4 1 Dicot 0.5 4 5.0 4 1b 2 1 Shrub 1 2 Shrub 1.1 Dicot Shrub 1 6.0 Dicot 2.5 0.5 1 5.0 0.0 2.1 1 1.4 Shrub 0.0 Shrub 1.5 Dicot 1.0 1 1b Dicot 1 1b 1 Dicot 4 1 1 Perennial forb Dicot Shrub 1 1 Annual Dicot forb Dicot Dicot 1 Perennial forb Perennial forb Annual forb Perennial forb Dicot Dicot Dicot Perennial Dicot forb Perennial forb Perennial forb Perennial forb FabaceaeFabaceaeFabaceaeFabaceae WIFabaceae SEFabaceae SIFabaceae 1.5 SIFabaceae 6.1 SEFabaceae 3 BE/SI 3.0 24.5 WI/IN 3.1 4.4 15 6.1 WI/IN 1.3 15.25 WI/IN 4 2 21.75 24.5 1.3 1.0 2.5 5 5 6.0 1.5 5 4 4 3.1 0.0 0.7 3.0 3.0 2 6.0 0.0 0.5 6.0 2.9 3.0 3 2.4 3 1.0 1b 0.0 1.3 1.3 1.1 1.4 1.0 0.0 1.0 4 1.0 0.6 1b 1.0 1 1b 1 0.3 1 0.5 1 1 1 1 1 1 1 1 Dicot 1 Dicot 1 Dicot Dicot Dicot 1 Dicot Perennial forb Dicot Perennial forb Perennial forb Perennial Perennial forb forb Annual Dicot forb Annual forb Annual forb Fabaceae WI 1.8 7.2 4 0.6 2.8 1.4 1 1 1 Dicot Perennial forb FabaceaeFabaceaeFabaceae BE/SI 4.0 WI/IN WI/IN 12 1.4 1.3 7 4 3 4.0 5 3 2.0 1.0 1.2 1.0 1.1 0.6 0.5 0.3 1 1 1 1 Dicot 1 Dicot Perennial forb Annual forb Fabaceae BE 5.3 21 4 6.0 1.5 0.8 1 1 Dicot Perennial forb ) Fabaceae SI 3.0 6 2 3.0 0.0 0.0 1b 1 Dicot Annual forb . elmeri ssp. ssp. var. claranus vestitus ( var. var. var. var. junceus dentata var. 1 ONTINUED var. ssp. Taxon 1. C raichei ssp. pulchella viscida jepsonianus siskiyouensis gracilentum oreganum PPENDIX A Arctostaphylos klamathensis Arctostaphylos nortensis Arctostaphylos obispoensis Arctostaphylos stanfordiana Arctostaphylos viscida Arctostaphylos viscida Pyrola picta Vaccinium coccineum Astragalus breweri Astragalus clarianus Astragalus curtipes Astragalus macrodon Astragalus rattanii Astragalus whitneyi Hoita strobilina Lathyrus bi¯orus Lathyrus delnorticus Lathyrus vestitus Astragalus clevelandii Lupinus constancei Lupinus lapidicola Lupinus onustus Lupinus spectabilis Pediomelum californicum Trifolium amoenum Trifolium fucatum Trifolium gracilentum Lotus junceus Trifolium microcephalum Trifolium willdenovii Trifolium longipes Trifolium longipes 244 MADRONÄ O [Vol. 52 11 (rhiz.) (rhiz.) (rhiz.) (rhiz.) Lifeform 10 Tax. Cat. Dicot Shrub Dicot Annual forb Dicot Perennial forb Dicot Perennial forb 9 1111 1111 1111 Geog. Dist. 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Fagaceae SI 2.5 12.25 5 1.0 2.5 1.1 1 1 Dicot Shrub Fagaceae SE 5.8 40.5 7 6.0 0.8 0.3 FagaceaeGarryaceaeGarryaceaeGentianaceae SI SEGentianaceae BEGentianaceae SEHydrophyllaceae SE WI 5.8 2.5 WI 5.0Hydrophyllaceae 5.5 29 12.25 BE/SI 5.8 30 1.8 11Hydrophyllaceae 1.5 4.3 5 17.5 SEHydrophyllaceae 7 5 WI/INHydrophyllaceae 3 17 6 2.0 SEHydrophyllaceae 2 3 1.4 5.5 SE 6.0Hydrophyllaceae 2.2 5.5 WI/INHydrophyllaceae 4 5.5 11 0.4 2 6.0 4.25 4 1.0 5.5 WIHydrophyllaceae 1.3 1.1 0.7 6.1 0.2 WI 0.3 2.0 33 1.5 4.5 3 0.5 0.5 0.2 30.5 2.1 2 0.5 2.3 2.1 1.7 1.0 2.1 1 0.3 1.5 0.9 6 5.5 7 5 1 2 1.4 1 6.25 1 0.7 1 1 0.8 6.0 6.0 1.1 1 0.5 3 1 3 1 0.8 0.0 1 1.3 1 1 0.3 3.0 0.0 1.0 3.0 1 4 1.6 1.2 Dicot 1 1 1 0.9 0.7 Dicot 1 Dicot 1 1 1 Dicot Shrub Dicot 1 1 1 1 Shrub Shrub 1 Dicot 1 Dicot Annual Perennial forb forb Dicot 1 1 Annual Dicot 1 forb Perennial forb 1 Perennial forb Dicot Dicot Dicot Annual forb Dicot Annual Perennial forb forb Perennial forb Annual forb Hydrophyllaceae SEHydrophyllaceae WI 6.1 1.7 36.5 5 6 3 6.0 0.0 1.0 0.0 1.2 1b 0.7 1 Dicot Annual forb Iridaceae WI 1.5 3 2 1.5 2.1 1.5 1 1 Monocot Perennial forb Hydrophyllaceae BE/SIHydrophyllaceae BE/SIHydrophyllaceae 3.9 BE/SIHydrophyllaceae 4.2 WI/INIridaceae 27.5 4.0 12.5 1.1Iridaceae 12 7 2.1 3 SEIridaceae 4.0 3 SE 6.0 2 1.1 5.8 3.2 4.0 WI/IN 0.4Lamiaceae 1.1 11.5 5.8 1.8Lamiaceae 1.0 1b 1.1 1.3Lamiaceae 11.5 0.6 2Lamiaceae 1 SE 1.0 3.25 WI/IN 2 5.5 SI 3 1.3 6.1 BE/SI 0.7 1 5.5 30.5 1.0 3 3.5 0.5 3.4 0.7 1 0.9 3 10.5 16.75 0.5 5 1 0.5 3 4 1 5 3 Dicot 6.0 0.0 1 Dicot 0.0 3.0 3.0 1.7 1 1 0.0 Annual 2.1 2.5 forb Dicot 1.0 Dicot 1b 1 Annual 0.9 1.5 forb 1 4 4 Annual forb Annual forb Monocot 1 1 Monocot Perennial forb Monocot 1 Perennial forb 1 Perennial forb 1 1 1 Dicot Dicot Dicot Dicot Annual forb Annual Annual forb forb Annual forb var. var. . ssp. var. durata ssp. 1 var. ONTINUED ssp. Taxon 1. C echinoides penduli¯ora rosea imbricata purdyiformis obovata PPENDIX Quercus durata Lithocarpus densi¯orus Quercus vaccinifolia Garrya buxifolia Garrya congdonii Centaurium tricanthum Gentiana setigera Swertia fastigiata Emmenanthe penduli¯ora Emmenanthe penduli¯ora Phacelia breweri Phacelia californica Phacelia corymbosa Phacelia dalesiana Phacelia distans Phacelia divaricata Phacelia egena Phacelia greenei Phacelia imbricata Phacelia leonis Phacelia phacelioides Phacelia pringlei Phacelia purpusii Iris bracteata Iris innominata Iris macrosiphon Iris tenuissima Acanthomintha duttonii Acanthomintha ilicifolia Acanthomintha lanceolata Acanthomintha obovata A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 245 11 (bulb) (rhiz.) (rhiz.) (rhiz.) (carn.) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) Lifeform 10 Tax. Cat. Dicot Perennial forb Dicot Shrub Dicot Perennial forb Monocot Perennial forb 9 Geog. Dist. 1111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Lamiaceae SE 6.1 24.5 4 6.0 0.0 0.0 4 1 Dicot Perennial forb LamiaceaeLamiaceaeLamiaceae SILamiaceae SE BE 3.0 BE/SI 5.8 6 4.8 34.5 4.4 28.5 22 2 6 6 3.0 6.0 5 6.0 1.4 0.8 2.2 1.0 6.0 0.3 0.9 2.3 1b 1b 1.0 1 1 1 1 1 Dicot Dicot Dicot Annual forb Shrub Perennial forb LamiaceaeLamiaceaeLamiaceae SILamiaceae SE BE/SI 3.0 WI 6.1 4.3 18 30.5 17 1.6 6 5 9.5 4 3.0 6.0 6 4.5 1.7 0.0 0.7 0.0 2.1 1.5 1.0 1b 1.3 0.5 1 1 1 1 1 Dicot Dicot Dicot Perennial Perennial forb forb Perennial forb LamiaceaeLamiaceae WI WI 2.3 2.2 11.5 11 5 5 3.0 1.5 1.0 0.7 2.4 1.1 1 1 1 Dicot Perennial forb LamiaceaeLamiaceaeLentibulariaceae SE BE/SILiliaceae BE 4.0Liliaceae 6.2 16 5.4 18.5 WI 21.5 WI 4 3 1.5 4 4.5 6.0 2.3 4.5 6.0 2.4 0.0 11.25 1.5 1.2 0.0 3 0.8 5 2 2.0 4 2.0 0.9 1 1 2.2 0.5 1 1.0 1 1 1 1 1 1 Dicot Dicot Dicot Annual Perennial forb Annual forb forb Monocot Perennial forb Liliaceae WI/IN 1.1 4.5 4 1.0 0.6 0.3 1 1 1 Monocot Perennial forb Liliaceae BELiliaceae 5.4 BE/SI 21.5 4.0 4 12 6.0 3 1.5 0.8 4.0 4 1.0 0.6 1 Monocot 1 Perennial forb Monocot Perennial forb LiliaceaeLiliaceae SILiliaceae WI/INLiliaceae 2.6 1.3 SE 15.75 3.75 BE/SILiliaceae 6.0 6 4.2 3 18 2.5 38 SE 1.0 1.9 0.7 3 0.8 9 6.1 0.4 6.0 30.5 4.0 0.0 1.6 1 0.0 5 0.5 1 6.0 0.0 1 1 0.0 1 1 1 4 Monocot 1 1 Monocot Perennial forb 1 1 Perennial forb Monocot Monocot Perennial Perennial forb forb Monocot Perennial forb Liliaceae WI 2.0 4 2 2.0 0.0 0.0 1 1 Monocot Perennial forb . viridis purdyi ssp. bolanderi mirabile ssp. ssp. ssp. var. var. var. 1 var. ONTINUED Taxon 1. C benitensis douglasii macroceras sanbenitense PPENDIX Allium bolanderi A Monardella antonina Monardella douglasii Monardella follettii Monardella palmeri Monardella purpurea Monardella sheltonii Monardella stebbinsii Monardella viridis Salvia sonomensis Scutellaria antirrhinoides Stachys pycnantha Trichostema laxum Trichostema rubisepalum Pinguicula vulgaris Allium acuminatum Allium amplectens Allium bolanderi Allium cratericola Allium crispum Allium diabloense Allium falcifolium Allium ®mbriatum Allium hoffmanii Allium howellii 246 MADRONÄ O [Vol. 52 11 (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) Lifeform 10 Tax. Cat. 9 Geog. Dist. KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Liliaceae BE 4.5 13.5 3 4.0 0.6 0.3 4 1 Monocot Perennial forb LiliaceaeLiliaceae BE/SILiliaceae 3.8 BE/SILiliaceae 15.25 4.3 WI/INLiliaceae 13 1.3 4 WI/INLiliaceae 1.0 4 4.5 WILiliaceae 3 2.8 2 SELiliaceae 1.8 1.4 6.0 3 SILiliaceae 2.9 5.6 3.5 2 1.0 SILiliaceae 1.7 22.5 1.5 3.4 1.0 2 BELiliaceae 0.9 27 1.4 2.6 4 SILiliaceae 1.8 1.0 5.1 10.5 1 1.8 6.0 SELiliaceae 8 20.5 1 1 1.3 1.0 2.8 4 WI/INLiliaceae 1b 3.5 0.5 14 6.2 4 1.0 3.0 WI/INLiliaceae 2.2 4 Monocot 18.5 1 1.5 6.0 1.1 0.8 3 WILiliaceae Perennial 5 forb Liliaceae 0.7 2.0 4 3 4.5Liliaceae WI/IN Monocot 1 1.0 1 2.0Liliaceae 2.0 3 1.0 6.0 SE 1b Monocot 1 Perennial 1.8 forb 4 BE 4 1 0.0 1.0 0.8 2Liliaceae WI Perennial forb Monocot 1.3 0.0 SE 5.5 1.0 4 1 MonocotLiliaceae 4.9 1b 1.0 0.6 Perennial 2 27.5 1 forb 2.2 1 2 19.5 Perennial 1 0.5 WI forb 6.0 1 Monocot 11 1 2.0 1 5 1.0 18 1 1 WI Perennial 4 1.4 forb 1.5 1.4 Monocot 6.0 1.0 5 Monocot 5.0 1.0 2.0 4.5 1 3 1.3 Perennial Monocot forb Perennial 1.5 2.0 1 forb 1 0.6 4 Perennial 6.0 forb 0.8 3 0.8 1 1b 1 Monocot 0.0 1 1b 0.4 Monocot 1.0 1 0.0 2 1 Perennial forb 1.3 Monocot Perennial 1 forb 1 2.0 0.8 Monocot Perennial forb 1.4 Perennial forb 1.0 1 1 1 Monocot Monocot 1 1 Perennial Perennial Monocot forb forb 1 Monocot Perennial 1 forb Perennial forb Monocot Monocot 1 Perennial forb Monocot Perennial forb Perennial forb Monocot Perennial forb Liliaceae BE 5.4 37.5 7 6.0 1.0 0.4 1b 1 Monocot Perennial forb nanus rosea . var. congdonii sanbornii var. var. var. var. ssp. ssp. var. var. var. var. var. 1 var. ONTINUED Taxon 1. C clavatus lacunosum micranthum franciscanum coronaria ®mbriatus conspicuum californica leptandra PPENDIX Allium lacunosum Allium lacunosum Allium membranaceum Allium obtusum Allium peninsulare Allium sanbornii Allium sanbornii Allium serra Allium sharsmithiae Allium siskiyouense Allium tuolumnense Allium unifolium Brodiaea californica Brodiaea californica Brodiaea coronaria Brodiaea coronaria Brodiaea pallida Brodiaea purdyi Brodiaea stellaris Calochortus clavatus Calochortus coeruleus Calochortus elegans Allium jepsonii A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 247 11 (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) Lifeform 10 Tax. Cat. Monocot Perennial forb 9 Geog. Dist. 1111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Liliaceae WI 2.0 6 3 2.0 1.0 0.6 1 1 Monocot Perennial forb LiliaceaeLiliaceae SELiliaceae WILiliaceae 6.0 BELiliaceae 12 2.1 SELiliaceae 5.4 8.5 SELiliaceae 2 21.5 6.2 4 SI 6.0 18.5 6.1 4 WI 0.0 2.5 24.5 2.9 3 0.0 6.0 1.2Liliaceae 1.7 14.5 0.6 1.5 4 6.0Liliaceae 0.8 5 0.0 6.0 5 1 WILiliaceae 1b 0.0 0.0 1 3.0 BELiliaceae 1b 3 0.0 2.4 1.1 SELiliaceae 1b 1.0 5.2 0.5 9.5 SELiliaceae 1 1.2 4 26 6.1 4 0.7 SILiliaceae 1 1 30.5 5.5 1 Monocot 1 BE/SILiliaceae 1.8 5 16.5 Monocot 1 2.7 5 Perennial 2.8 4.3 Monocot forb BELiliaceae 6.0 1 Perennial forb 1.4 21.5 3 8 6.0 Perennial 1.1 Monocot BELiliaceae forb 4.7 0.0 0.5 Monocot 6.0 Perennial SILiliaceae forb 5 37.5 0.0 1b 4.5 3 1.2 Perennial forb WILiliaceae Monocot 1b 4.0 18 0.7 2.0 2.5 8 1 SI 0.4 Perennial 1b forb 2.1 2.3 5 0.2 4.5 WI/IN 4 1 1.2 4 7 1.4 3.0 1.0 1 4.5 0.5 2 15 1 1 1 2 1.7 1b 3 1 Monocot 1 2.5 0.9 1 1 1 5 Monocot Perennial 3.5 2.0 4 Monocot forb 2 2.5 2.5 Perennial 2.0 forb Perennial forb 1.0 1.5 Monocot 1.7 1.4 1 1b 0.8 Perennial forb 1.0 1 Monocot 1 Monocot Perennial forb Perennial Monocot forb Perennial forb Monocot Perennial forb Monocot Monocot Perennial 1 forb Perennial 1 forb Monocot Monocot Perennial forb Perennial forb Liliaceae WI/IN 1.0 3 3 0.0 1.7 1.0 1b 1 Monocot Perennial forb var. . vestus var. var. var. 1 ONTINUED Taxon 1. C minus var. reductum citrinum roderickii PPENDIX Calochortus vestae Calochortus weedii A Calochortus greenei Calochortus nudus Calochortus obispoensis Calochortus raichei Calochortus tiburonensis Calochortus umbellatus Calochortus uni¯orus Chlorogalum angustifolium Chlorogalum grandi¯orum Chlorogalum pomeridianum Chlorogalum purpureum Erythronium californicum Erythronium citrinum Erythronium citrinum Erythronium helenae Erythronium hendersonii Erythronium howellii Erythronium multiscapoideum Erythronium purpurascens 248 MADRONÄ O [Vol. 52 11 (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) (bulb) Lifeform 10 Tax. Cat. Monocot Perennial forb Monocot Perennial forb 9 1111 1111 Geog. Dist. KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff LiliaceaeLiliaceae WILiliaceaeLiliaceae SE 2.3Liliaceae BE/SI 13.5 WI 6.1Liliaceae 4.3 24.5 WILiliaceae 6 17.25 1.8 BE 4 2.0 4 2.4 7 0.4 SI 6.0 5.5 4.5 9.5 0.2 0.0 2.7 4 31.5 3 2.7 0.0 4 1.4 1.5 1b 7 8 2.5 1.1 4.0 1.5 0.6 1 3 0.7 1.8 1b 1 0.7 1 2.0 4 1 3.1 1 1.8 1 Monocot 1 1 1 Monocot 1 Perennial forb Monocot Perennial forb Perennial forb Monocot 1 1 Perennial forb Monocot Monocot Perennial forb Perennial forb Monocot Perennial forb Liliaceae BE 5.4 21.5 4 6.0 1.5 0.8 1b 1 Monocot Perennial forb Liliaceae WI 2.3 9 4 2.5 1.7 0.9 1 1 1 Monocot Perennial forb LiliaceaeLiliaceae WI SI 2.0 6 2.7 13.25 3 5 2.0 2.0 0.0 1.6 0.0 0.7 4 1 1 1 1 Monocot Perennial forb Liliaceae SI 2.5 5 2 2.5 3.5 2.5 1 Monocot Perennial forb LiliaceaeLiliaceae SELiliaceae SILiliaceae 6.2 SELiliaceae 18.5 3.4 SELiliaceae 17 6.0 3 SILiliaceae 18 6.2 6.0 WILiliaceae 5 18.5 0.0 2.5 BE/SI 3 3.0 0.0 10 2.0 3 3.5 1b 1.5 WI 6.0 9.75 10.5 0.7 6.0 0.0 4 2.0 0.0 0.0 5 3 2.0 0.0 6 2.0 1 4 1.9 3.0 1.4 1 1.0 1 1 2.5 1 0.6 3 1 1.5 4 2.0 4 Monocot 1 1.0 1 1 1 Perennial 1 forb 0.6 1 Monocot 1 Perennial forb Monocot Monocot Perennial forb Perennial forb Monocot Monocot Perennial Monocot Perennial forb forb Perennial forb Liliaceae SI 2.7 8 3 3.0 0.6 0.3 1 1 1 Monocot Perennial forb LiliaceaeLiliaceae SI SI 2.7 3.3 8 13 3 4 3.0 3.5 0.6 1.7 0.3 0.9 1 1 1 1 1 Monocot Monocot Perennial Perennial forb forb ssp. . recurva ineziana bi¯ora af®nis var. var. var. var. var. 1 ONTINUED Taxon 1. C coccinea purpurascens PPENDIX Fritillaria eastwoodiae Fritillaria falcata Fritillaria glauca Fritillaria liliacea Fritillaria pluri¯ora Fritillaria purdyi Fritillaria recurva Fritillaria recurva Fritillaria bi¯ora Fritillaria bi¯ora Fritillaria af®nis Fritillaria agrestis Erythronium tuolumnense Fritillaria viridea Hastingsia alba Hastingsia serpentinicola Lilium bolanderi Lilium kelloggii Lilium rubescens Lilium washingtonianum Muilla maritima Odontostomum hartwegii Triteleia bridgesii A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 249 11 (rhiz.) (bulb) (bulb) Lifeform 10 Tax. Cat. Dicot Annual forb Dicot Annual forb Dicot Annual forb Dicot Perennial forb Dicot Annual forb Monocot Perennial forb Monocot Perennial forb 9 1111 1111 Geog. Dist. 11111 11111 11111 1111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff OnagraceaeOnagraceae BE WI 5.0 2.0 25 6 5 3 5.0 2.0 1.0 0.4 1.0 0.6 4 1 Dicot Annual forb Onagraceae BE/SI 3.8 23 6 4.0 2.2 0.9 1 1 Dicot Perennial forb Onagraceae WI 2.2 6.5 3 2.0 1.0 0.6 1b 1 Dicot Annual forb OnagraceaeOnagraceaeOnagraceae WI/IN BE/SI 1.4 SE 3.8 2.75 11.5 6.1 2 24.5 3 1.4 3.0 4 0.9 2.1 0.6 6.0 1.2 0.0 4 0.0 1b 1 1 1 1 1 1 Dicot Dicot Dicot Annual forb Annual forb Annual forb Liliaceae BE/SI 3.8 23 6 4.0 0.8 0.3 4 1 1 1 Monocot Perennial forb LiliaceaeLinaceaeLinaceae WILinaceaeLinaceae SE 1.6 SE SI 4.75 SI 5.7 6.2 28.5 3 2.5 18.5 2.8 10 2.0 5 3 8.5 0.7 6.0 0.4 4 6.0 0.9 3 0.0 2.5 0.4 3.0 0.0 1b 1.5 1 0.6 1b 0.7 1 0.3 1b 1 1 1 1 1 Monocot Perennial forb Dicot Dicot Dicot Annual forb Annual forb Annual forb LinaceaeLinaceaeLinaceaeLinaceae WILinaceae SELinaceae SE 2.0 SE SE 6.1 WI 8 6.2 24.5 6.0 18.5 6.1 18 2.4 4 4 24.5 3 11.75 6.0 2.0 3 4 6.0 5 0.0 1.8 0.0 6.0 0.0 6.0 0.9 3.0 0.0 0.0 1b 0.0 1.0 1b 0.0 0.0 0.4 1b 1 1 1 1 1 1 1 1 1 Dicot Dicot Dicot Annual Dicot forb Dicot Annual forb Annual forb Annual Annual forb forb LinaceaeLinaceaeLinaceaeLinaceae SE BE SE WI/IN 6.2 4.7 1.3 18.5 5.8 14 34.5 4 3 3 6 6.0 3 0.0 6.0 6.0 1.0 0.0 2.3 0.8 1.5 1b 1.3 0.3 0.9 1b 1 1 1 1 Dicot Dicot Dicot Annual forb Annual Annual forb forb Malvaceae SI 2.6 13 5 3.0 2.3 1.0 MalvaceaeMalvaceaeMalvaceae WIMalvaceae SEOnagraceaeOnagraceae 1.6 SEOnagraceae SI 5.6 SE 4.75 SI 22.5 6.3 WI 3 3.0 12.5 6.1 4 3.0 24.5 2.0 6 2.3 2 6.0 0.7 9 7 4 1.0 0.4 6.0 2 0.5 0.0 6.0 3 3.0 1b 3 0.0 0.0 2.8 3.0 1b 0.0 2.0 2.0 3.0 1b 1 0.6 1b 1.7 0.3 1 1 1 1 Dicot 1 Dicot 1 Dicot Annual forb 1 Perennial forb Dicot Dicot Perennial forb 1 1 Dicot Annual forb Annual Dicot forb Annual forb Annual forb LiliaceaeLiliaceaeLiliaceae SILiliaceae BE BE 3.3 4.5 BE/SI 4.5 10 22.5 3.8 13.5 19 5 3 3 4.0 3.0 5 6.0 1.5 2.5 2.9 3.0 0.7 1.5 1.7 2.2 4 4 1b 1.0 1 1 1 Monocot Monocot Monocot Perennial Perennial forb forb Perennial forb Liliaceae WI 1.6 8 5 1.0 0.9 0.4 . viridis var. cookii tracyi albicaulis crocea modesta biloba ssp. ssp. ssp. 1 var. var. ssp. ssp. ONTINUED ssp. Taxon 1. C fontanus anomala PPENDIX Epilobium minutum Epilobium oreganum Clarkia gracilis Clarkia breweri Clarkia franciscana Clarkia gracilis Zigadenus micranthus Zigadenus paniculatus Hesperolinon adenophyllum Hesperolinon bicarpellatum Hesperolinon breweri Hesperolinon californicum Hesperolinon clevelandii Hesperolinon congestum Hesperolinon didymocarpum Hesperolinon disjunctum Hesperolinon drymarioides Hesperolinon micranthum Hesperolinon serpentinum Hesperolinon spergulinum Hesperolinon tehamense Linum lewisii Sidalcea diploscypha Sidalcea hartwegii Sidalcea hickmanii Sidalcea hickmanii Sidalcea keckii Camissonia benitensis Camissonia lacustris Clarkia arcuata Clarkia biloba Triteleia crocea Triteleia ixioides Triteleia peduncularis A Triteleia crocea Xerophyllum tenax 250 MADRONÄ O [Vol. 52 11 gram. (cesp.) gram. (rhiz.) gram. (rhiz.) gram. (cesp.) gram. (cesp.) gram. (cesp.) (bulb) (bulb) (bulb) (paras.) Lifeform 10 Tax. Cat. DicotGymnosp. Annual Tree forb Gymnosp.Dicot Tree Annual forb Monocot Perennial Dicot Perennial forb 9 Geog. Dist. 11111 11111 11111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff PapaveraceaePapaveraceaePapaveraceae SEPapaveraceae WI SI WI 5.6 2.2 22.5 2.6 1.7 6.5 7.75 4 5 3 6.0 3 3.0 1.0 3 1.0 1.4 0.5 3.1 2.0 0.8 4 1.8 0.6 4 1 0.3 1 1 1 Dicot Dicot Dicot Perennial forb Perennial forb Annual forb PoaceaePoaceae WI 1.7 SE 5 6.1 24.5 3 4 2.0 1.5 6.0 0.9 0.0 0.0 4 1 1 1 Monocot Monocot Perennial Perennial Poaceae WI/IN 1.0 2 2 1.0 1.4 1.0 1 Monocot Perennial PinaceaePinaceaePinaceaePinaceae WIPinaceae SIPinaceae BE/SIPlantaginaceae 2.2 4.3Poaceae WI/IN 2.5 WI/IN SI 6.5 26 1.3 WI/IN 12.6 1.0 1.4 4 3 BE 2.7 3 5 6 4.25Poaceae 3.0 8 3.0 4.8 4.0 3 1.4 3 3 2.4 1.5 14.5Poaceae 0.8 1.0Poaceae 3 1.0 1.1 WI/IN 1.0 0.6 1.5 3 1.0 1.4 1.1 3.0 0.9 0.6 WI/IN 0.8 1 0.6 6.0 2.1 WI 1 1.1 0.3 2.3 2 1.3 4.25 1.7 3 65 1.1 4 1 1.3 1 1 1 1.1 1.0 3 1 1 1.1 Gymnosp. Gymnosp. 2.0 0.5 Tree 1 Tree 0.6 Gymnosp. 1 Tree 0.3 Gymnosp. Tree Monocot 1 Perennial 1 1 1 Monocot Monocot Perennial Annual gram. OrchidaceaeOrchidaceae SIOrobanchaceae WI/IN SI 2.5Papaveraceae 1.2 12.25 WI/IN 3.5 3.4 1.1 5 13.5 3 3.25 2.0 4 1.0 1.6 3 1.0 0.7 3.0 0.6 1.0 4 1.3 4 0.9 0.6 1 0.5 4 1 1 1 1 1 1 1 Monocot Monocot Perennial forb Perennial forb Dicot Perennial forb OnagraceaeOnagraceaeOrchidaceae BE SE BE 5.1 5.5 20.5 4.5 38.5 40.5 4 7 9 6.0 6.0 2.0 4.0 1.0 1.0 1.3 0.4 4 0.4 1b 4 1 1 1 1 1 1 Dicot Monocot Dicot Perennial forb Perennial forb Perennial forb dorei . var. howellii oregana var. ssp. ssp. ssp. 1 ONTINUED Taxon 1. C pubescens balfouriana PPENDIX Dicentra pauci¯ora Eschscholzia hypecoides Platystemon californicus Calamagrostis foliosa Calamagrostis ophitidis Picea breweriana Pinus attenuata Pinus balfouriana Pinus coulteri Pinus jeffreyi Pinus sabiniana Plantago erecta Achnatherum lemmonii Achnatherum nelsonii Achnatherum stillmanii Agrostis microphylla Bromus laevipes Cypripedium fasciculatum Piperia candida Orobanche valida Dicentra chrysantha Epilobium rigidum Epilobium siskiyouense Cypripedium californicum Dicentra formosa A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 251 11 gram. (cesp.) gram. (cesp.) gram. (rhiz.) gram. (rhiz.) gram. (cesp.) Perennial forb Perennial forb gram. (cesp.) gram. (cesp.) gram. (cesp.) gram. (cesp.) gram. (cesp.) Lifeform 10 Tax. Cat. MonocotMonocot Annual gram. Annual gram. Monocot Perennial Monocot Perennial Monocot Perennial Monocot Perennial Monocot Perennial Monocot Perennial 9 Geog. Dist. 11111 11111 11111 11111 11111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff PoaceaePoaceae WI/IN 1.2Poaceae BE 6Poaceae 5.4 WI 5 21.5Poaceae 1.8 SI 1.0 4 0.4 3.5 WI 3.3 0.2 5.5 2 13 1.0 1.7 0.5 1.5 1 4 5.1 4 2.1 1 1.5 1 3.0 3 4 1.3 1 1.0 0.6 1 1 2.0 Monocot 1.2 Perennial Monocot Perennial 1 Monocot 1 1 Perennial Monocot Perennial PoaceaePolemoniaceae WI WI 2.3 1.6 9.1 4.75 4 3 2.0 1.0 2.0 1.2 1.0 0.7 1 1 Dicot Annual, PolemoniaceaePolemoniaceae SE WI 5.6Polemoniaceae 1.8 33.5 BE/SI 7.1 3.8 6 19 4 6.0 2.0 1.2 5 1.5 0.5 0.7 4 3.0 2.2 1.0 1 1 4 1 1 1 1 Dicot Dicot Annual forb Annual forb Dicot Annual, Poaceae SI 3.1 9.25 3 3.0 2.9 1.7 Poaceae WI/IN 1.3 5.25 4 1.0 1.2 0.6 Poaceae WI 2.4 11.75 5 2.0 1.6 0.7 Poaceae WI 1.6 3.1 2 1.6 2.1 1.5 Poaceae SI 3.3 13 4 3.0 2.2 1.1 Poaceae WI 1.5 3 2 1.5 2.1 1.5 ssp. . var. ssp. ssp. var. capitata pinnatisecta 1 ONTINUED ssp. ssp. Taxon 1. C microstachys californicum trachycaulus californica inexpansa PPENDIX Melica geyeri Poa piperi Poa rhizomata Poa tenerrima Scribneria bolanderi Vulpia microstachys Collomia diversifolia Collomia tinctoria Gilia capitata Gilia sinistra Hordeum brachyantherum Festuca idahoensis Festuca californica Elymus trachycaulus Danthonia californica A Calamagrostis stricta 252 MADRONÄ O [Vol. 52 11 forb, Shrub Perennial forb (rhiz.) Lifeform 10 Tax. Cat. Dicot Annual forb Dicot Annual forb DicotDicot Annual forb Annual forb 9 1111 Geog. Dist. 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Polemoniaceae BEPolemoniaceaePolemoniaceae SIPolemoniaceae 5.3 SEPolemoniaceae SE 31.5 WI 2.8 5.9 14 5.6 6 23.5 2.0 22.5 6.0 6 5 4 1.3 4 0.5 3.0 6.0 3 5.5 1b 2.4 0.5 0.6 1.1 0.3 2.0 0.3 4 1.0 1 4 0.6 1 1 Dicot 1 Perennial forb Dicot Dicot Dicot Annual forb Annual forb Annual forb PolemoniaceaePolemoniaceae SEPolemoniaceae SEPolygalaceae WI 6.0Polygonaceae WI 6.2Polygonaceae 18 1.6Polygonaceae BE 18.5 BE 2.3 8.1 SI 3 3 5.4Polygonaceae 9 4.9Polygonaceae 5 21.5 6.0Polygonaceae 6.0 BE 2.7 24.5Polygonaceae SE 0.0 2.0 0.0 4 10.75 SI 4 0.0 5.3 1.5Polygonaceae 0.0 WI 5 2.0Polygonaceae 6.1 4 1b 5.5 0.7 1b 16Polygonaceae BE 3.1 6.0 1.3 30.5 1b 1.7 1.0Polygonaceae SI 2.5 1 0.6 12.5 1.6Polygonaceae SI 0.5 1 2.0 5.1 3Polygonaceae SI 5.1 1 5 0.7 1b SI 1.0 3.0 35.5 1 4 4Polygonaceae SE 6.0 3.0 6.0 3 12Polygonaceae 1 3.2 1.2 3.0Polygonaceae SI 0.0 7 3.3 6 2.0Polygonaceae SE 6.2 0.7 2.6 9.5 0.0 13 SE 4 6.0 1.5 4 18.5 1.3 1b SE Dicot 1 3.3 2 Dicot 0.9 1.7 6.1 3 4 3.0 1 Dicot 1 13 6.1 4 3 0.7 24.5 3.0 1.6 5.5 1 3.0 Annual forb 1 36.5 4 Perennial forb 0.8 3.5 6.0 1.4 1 27.5 3.0 Perennial Dicot 1 forb Dicot 4 4 1.0 3.2 0.0 1 1 1.7 6 Dicot 1 5 3.5 1.6 6.0 0.0 1 Dicot 4 1 Perennial Annual 6.0 forb 1b 3.2 0.0 1 6.0 Annual 0.0 1 forb 1.6 0.0 Annual, 1 0.9 Dicot Dicot 0.0 1 1b 1 0.4 Dicot 4 3 Dicot Perennial Annual 1 forb forb 1 Dicot Annual 1 forb 1 Perennial forb Dicot 1 Shrub 1 Dicot Dicot 1 Dicot Dicot Annual forb Perennial Annual forb forb Dicot Perennial forb Perennial forb Dicot Dicot Perennial forb Perennial forb Annual forb PolemoniaceaePolemoniaceae SIPolemoniaceae SEPolemoniaceae WI/INPolemoniaceae SI 2.5 1.3 WI 5.8Polemoniaceae 7.5 17.5 2.5 WI 2.5 2.0 12.35 3 3 2 1.6 6 5 3.0 6.0 1.3 6.25 1.8 0.6 1.1 3.0 3 1.0 4 0.3 0.8 2.4 2.0 4 1.1 1.5 0.0 1.2 1 0.0 0.6 1 1 1 1 1 1 1 1 Dicot Dicot Dicot Annual Annual forb Annual forb forb Dicot Annual forb ) . var. howellii villosum cornuta ϭ var. ϭ ( ( var. ssp. sinistra var. ) ) 1 ONTINUED ssp. Taxon 1. C Leptosiphon n. s. h. ϭ compositum caninum Leptosiphon la. Leptosiphon li. ( PPENDIX Navarretia heterodoxa Navarretia jaredii Navarretia jepsonii Navarretia pubescens Navarretia rosulata Phlox hirsuta Polemonium chartaceum Polygala cornuta Chorizanthe breweri Chorizanthe palmeri Chorizanthe uniaristata Chorizanthe ventricosa Eriogonum alpinum Eriogonum argillosum Eriogonum compositum Eriogonum congdonii Eriogonum covilleanum Eriogonum dasyanthemum Eriogonum diclinum Eriogonum elatum Eriogonum hirtellum Eriogonum hirti¯orum Eriogonum kelloggii Eriogonum libertini Eriogonum luteolum Linanthus ambiguus Linanthus bolanderi Linanthus dichotomus Linanthus lini¯orus Gilia sinistra Linanthus latisectus Linanthus nuttallii A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 253 11 (rhiz.) Lifeform 10 Tax. Cat. Dicot Annual forb Dicot Annual forb 9 Geog. Dist. 1111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Portulacaceae BE/SI 3.6 18 5 3.0 1.3 0.6 Portulacaceae SI 3.1 15.5 5 3.0 0.2 0.1 1 1 1 Dicot Annual forb Portulacaceae SI 3.4 24 7 3.0 1.1 0.4 PolygonaceaePolygonaceae WIPolygonaceae SI 1.5PortulacaceaePortulacaceae SI BE 4.5 3.0 WI/IN 18.25 3.1 1.3 3 4.6 12.25 6 27.5 4 2.0 4 2.5 0.9 6 2.1 0.5 3 2.5 0.9 4.0 2.2 1.0 1.2 1.1 1.5 1 0.5 4 0.9 1 4 1 1 1 1 1 1 1 1 1 Dicot Dicot 1 Annual Dicot forb Annual forb Dicot Dicot Annual forb Perennial Annual forb forb PolygonaceaePolygonaceae SEPolygonaceae SEPolygonaceae 5.9 SE 29.5Polygonaceae SI 6.0Polygonaceae 24 BE 6.2 5Polygonaceae SI 3.3 18.5 6.0 5.3 10Polygonaceae BE/SI 4 0.4 3.0 3 26.5 3.5Polygonaceae SI 6.0 0.2 12 3 21 6.0 0.0Polygonaceae 4 BE 5 0.0 0.0 3.3 3.0Polygonaceae BE/SI 4 1 6.0 0.0 6 4.5 10Polygonaceae 2.5 4.2 1 1.3 4 WI/IN 3.5 1.5 27.25 3.0 WI/IN 0.6 21 1 1.1 1.4 3 1 1.2 1.0 6 4 1 0.7 2.25 1 0.5 3.0 3 5 5.0 4 2 2.5 2.1 4.0 1 Dicot 1.5 1 0.8 1.1 3 1.3 1 1 4 1.2 1 0.6 Dicot 1.0 Perennial 1 forb 0.9 1 Dicot 1 1 0.0 1 1 Perennial 1 0.0 forb Dicot Dicot 1 Perennial forb 1 Dicot Dicot Shrub Annual 1 forb Perennial forb Perennial forb Dicot Dicot 1 1 Dicot Perennial forb Dicot Perennial forb 1 Perennial Dicot forb Perennial forb Annual forb PolygonaceaePolygonaceae WIPolygonaceae WI 1.5PolygonaceaePolygonaceae SE 2.0 WI/IN 4.5 BE 1.0 6.2 6 3 18.5 3 5.4 1.0 3 32.5 3 0.6 3 2.0 0.3 6 6.0 0.0 0.0 4 0.0 6.0 0.0 1.7 0.0 1.2 1.0 2 0.5 4 1 1 1 1 1 1 Dicot 1 Dicot 1 Perennial forb Dicot Dicot Perennial Dicot forb Perennial forb Perennial forb Perennial forb PolygonaceaePolygonaceae BE/SI 3.8 SE 15 6.2 18.5 4 3.0 3 1.5 6.0 0.8 0.0 0.0 1b 1 1 1 1 1 Dicot Dicot Annual forb Perennial forb . var. var. var. var. var. glauca exigua ssp. ssp. var. var. var. var. var. var. ssp. ssp. 1 ONTINUED Taxon 1. C majus spergulariiforme proliferum hooveri argus bahiiforme goodmanii humistratum speciosum greenei indictum oblongifolium luteolum PPENDIX Claytonia gypsophiloides Claytonia exigua Polygonum douglasii Systenotheca vortriedei Calyptridium quadripetalum Calyptridium umbellatum Claytonia exigua Eriogonum strictum Eriogonum ternatum Eriogonum trichopes Eriogonum tripodum Eriogonum umbellatum Eriogonum umbellatum Eriogonum umbellatum Eriogonum umbellatum Eriogonum umbellatum Eriogonum ursinum Eriogonum vimineum Polygonum douglasii Eriogonum nudum Eriogonum pendulum Eriogonum pyrolifolium Eriogonum siskiyouense Eriogonum strictum A Eriogonum luteolum Eriogonum nervulosum Eriogonum nudum 254 MADRONÄ O [Vol. 52 11 (rhiz.) Lifeform 10 Tax. Cat. Dicot Perennial forb Pteridoph. Perennial forb Pteridoph. Perennial forb Dicot Shrub 9 Geog. Dist. 11111 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff PortulacaceaePortulacaceaePortulacaceae BE/SI WI/IN 4.4Portulacaceae WI 1.0Portulacaceae 21.75Portulacaceae WI/IN 6 2.0Portulacaceae WI/IN 5 1.0Portulacaceae SI 1.3 6Portulacaceae WI 5.0 4 BE 6 4 WI/IN 2.2 3.0 1.7 1.0 3 1.0 1.4 5.3 4 6 1.1 5 3 3.0 21 7 0.4 0.5 1.7 1.0 1b 1.4 2 1 3 1.0 1.5 4 5 0.7 1 1 3.0 0.9 2.0 6.0 1.0 0.0 1.5 1.5 1 1.1 0.0 1 0.9 0.8 0.5 1 Dicot 1 1 1 1 Dicot 1 1 Annual forb Perennial forb Dicot Dicot Dicot 1 1 Perennial forb Dicot Perennial Dicot forb Perennial forb Dicot Perennial forb Perennial forb Perennial forb RhamnaceaeRhamnaceae WIRhamnaceaeRhamnaceae SE 1.5 WI WI/IN 3 5.7 1.3 1.7 28.5 4 5 2 5 1.5 3 3 6.0 2.1 0.9 2.0 1.5 2.0 0.4 1.2 1.5 0.7 0.9 1b 1b 1 1 1 1 Dicot Dicot 1 Dicot Dicot Shrub Shrub Shrub Shrub PortulacaceaePortulacaceae BEPrimulaceae WIPteridaceae SI 4.7Pteridaceae 1.7 14 WIPteridaceae 3.0 BE 5PteridaceaeRanunculaceae 2.4 3 9 SIRanunculaceae WI 5.3 WIRanunculaceae 3 11.75 BE/SI 6.0 26.5 SIRanunculaceae 3.4 3 4.2 2.0 2.3 2.3 5 1.5Ranunculaceae WI/IN 31 1.3 1.5 25 5 3.0 6.75 2.0 BE/SI 2.7 4.5Ranunculaceae 1.4 1b 0.9 0.0Rhamnaceae 6.0 1.2 3.7 SE 3 8 9Rhamnaceae 0.0 4.1 6 3 0.5 1.1 11 WI/IN 2.0 0.5Rhamnaceae 1 3.0 3.5 WI 5.7 1 2.0 3Rhamnaceae 1.3 3 1.6 4 1.2 1.5 1 0.9Rhamnaceae 28.5 3 WI 0.9 1.0 0.4 Rhamnaceae 2.5 0.6 3.0 1.5 0.5 SERhamnaceae 4.0 1.5 BE/SI 5 0.6 2.0 6.1 SE 2 2.5 0.9 0.3 4.0 1 6.1 SI 6.0 1 1.5 1 Dicot 4 1 1 1 12 1.3 4 24.5 0.9 6.0 Dicot 1 1 1 1.1 1 3.3 0.4 18 1.5 1 Perennial 1 4 2 forb 0.8 Dicot 1 3 4 1.3 6.5 Perennial 1 forb 1b 1 Pteridoph. 6.0 2.0 0.6 3.0 3 1 Perennial Perennial forb forb 1 0.0 1.4 2 1 1.7 6.0 1 0.0 Dicot 1.0 Dicot Pteridoph. 1 1.0 3.0 0.0 1b 1b Perennial 1 1 forb 4.2 0.0 Perennial Dicot Dicot Perennial forb forb 3.0 Dicot 1b 1 Perennial Perennial forb forb Dicot 1 Dicot Perennial forb 1 1 1 Perennial 1 forb Shrub 1 Dicot Dicot Dicot Shrub Shrub Dicot Dicot Shrub Shrub Shrub ssp. . medius ssp. heckneri howellii var. var. var. ssp. var. var. var. 1 ONTINUED Taxon 1. C cotyledon roweanus patulum hesperium eastwoodiae cuneatus PPENDIX Claytonia saxosa Lewisia cantelovii Lewisia cotyledon Lewisia cotyledon Lewisia cotyledon Lewisia leana Lewisia nevadensis Lewisia oppositifolia Lewisia rediviva Ceanothus pumilus Ceanothus roderickii Ceanothus sonomensis Lewisia stebbinsii Lewisia triphylla Dodecatheon clevelandii Adiantum aleuticum Aspidotis carlotta-halliae Aspidotis densa Pellaea brachyptera Anemone drummondii Aquilegia eximia Delphinium hesperium Delphinium nuttallianum Delphinium parryi Delphinium uliginosum Ceanothus confusus Ceanothus cuneatus Ceanothus divergens Ceanothus ferrisae Ceanothus foliosus Ceanothus jepsonii Ceanothus masonii Ceanothus papillosus A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 255 11 (rhiz.) (carn.) Lifeform 10 Tax. Cat. Dicot Shrub Dicot Shrub Dicot Shrub Dicot Tree, shrub 9 1111 Geog. Dist. 11111 11111 1111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff RhamnaceaeRhamnaceae SERhamnaceae BE 6.0 WI 24 4.8 19 1.5 4 6 4 6.0 0.0 6.0 4 0.0 2.5 0.8 1.3 1.7 1 0.8 1 1 1 1 Dicot Dicot Shrub Shrub Rosaceae WI/IN 1.3 5.2 4 1.1 1.4 0.7 Rosaceae WI/IN 1.0 3 3 1.0 1.0 0.6 RosaceaeRosaceaeRosaceae BE/SIRosaceae 3.8 BE/SIRosaceae SERosaceae 3.8 7.5 SIRosaceaeRosaceae 15 WI 5.6 2 BERubiaceae 3.0 22.5 SI 3.5 4 1.6 BE/SIRubiaceae 9 5.4 0.7 4 SI 4.2 3.0 3.25Rubiaceae 3.1 32.5 0.5 12.5 6.0 1.5 SE 3 12.5Rubiaceae 2 2 3.3 0.8 1.0Rubiaceae 6 SI 2.0 3 1.6 0.5 10 1 4 5.5Rubiaceae 6.0 1.7 BE 1.9 4Rubiaceae 3.0 27.5 WI/IN 3.0 3.2 1.0 1.0Rubiaceae 1 1.4 1.7 3 1.4 0.0 1 0.4 5.1 16 WI/IN 1 5Salicaceae 1.0 SE 0.0 1b 3.0 20.5Salicaceae 1.0 2.75 SE 2 6.0Salicaceae 1b 1 2.5 1 1 5 2 SE 2 6.1 0.9 1 4 1.5 1 1 1 SE 5.9 Dicot 3.0 0.4 24.5 WI 1 1.4 5.0 29.5 6.0 1.9 2 0.9 Dicot 0.8 6.2 4 0.9 30 Perennial 1.6 forb 0.6 Dicot 1.0 5 0.4 18.5 1 1 1 6.0 1 Perennial 1.4 4 4.75 forb Dicot 1 6.0 Dicot 5 0.0 3 1.0 Perennial Dicot forb 0.4 3 0.0 Dicot 1b Dicot 6.0 Perennial 6.0 0.2 forb Perennial 1 1b forb 1.0 0.0 Perennial 1b 0.0 forb 1 1 Perennial 1.2 forb Perennial 0.0 1 0.0 forb 1 Dicot 1 0.7 1 Dicot 4 1 Perennial forb 1 Dicot 1 Perennial Dicot forb 1 Dicot 1 Perennial forb 1 Dicot Perennial forb Perennial Dicot forb 1 Dicot Perennial forb Perennial forb Dicot Perennial Dicot forb Shrub Shrub Sarraceniaceae BE/SISaxifragaceaeSaxifragaceae 4.1 WIScrophulariaceae WI BE/SIScrophulariaceae 32.5 WI 3.8 2.0 2.2 8 1.6 7.5 6 11 4.0 3.1 1.4 2 3 5 0.5 2 3.5 4 2.0 2.0 2.1 1.6 0.0 0.8 1.5 1 2.1 0.0 0.4 4 1.5 1 1 1 1 1 Dicot 1 1 Perennial forb Dicot Dicot 1 Dicot Dicot Perennial Perennial forb forb Annual forb Annual forb . gatense ssp. ssp. ssp. ssp. ssp. var. var. ssp. ssp. ssp. ssp. 1 ONTINUED Taxon 1. C occidentalis crassifolia tomentella nemorosa ambiguum siskiyouense andrewsii intermedium ¯avescens scotticum PPENDIX A Rhamnus californica Rhamnus tomentella Rhamnus tomentella Adenostoma fasciculatum Holodiscus discolor Horkelia congesta Horkelia daucifolia Horkelia sericata Horkelia tridentata Ivesia gordonii Ivesia pickeringii Potentilla cristae Sanguisorba of®cinalis Galium ambiguum Galium ambiguum Galium andrewsii Galium andrewsii Galium andrewsii Galium clementis Galium hardhamiae Galium serpenticum Salix breweri Salix delnortensis Salix sitchensis Darlingtonia californica Parnassia californica Saxifraga howellii Antirrhinum cornutum Antirrhinum leptaleum 256 MADRONÄ O [Vol. 52 11 (hemipar.) forb, Shrub (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) (hemipar.) Lifeform 10 Tax. Cat. Dicot Annual forb Dicot Annual forb Dicot Annual forb Dicot Perennial Dicot Annual forb 9 1111 1111 Geog. Dist. 11111 11111 11111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Scrophulariaceae BE/SIScrophulariaceae SI 4.3 21.5 2.5 20 5 4.0 8 1.8 0.8 2.5 4 1.3 0.5 1 Dicot Annual forb Scrophulariaceae SI 2.5 10 4 2.5 0.6 0.3 1 1 Dicot Annual forb Scrophulariaceae SIScrophulariaceae WI 2.8Scrophulariaceae SE 8.5 2.3Scrophulariaceae BE 6.2 9Scrophulariaceae 3 SI 18.5 4.6Scrophulariaceae 3.0 SI 4 27.5 0.6Scrophulariaceae 3.3 3 WI 0.3 2.5 16.5Scrophulariaceae 3.2 6 6.0 SE 4 1.0 2.4 15.75Scrophulariaceae 0.0 4.5 5 0.5 BEScrophulariaceae 0.0 WI/IN 5.6 9.75 1.4Scrophulariaceae 5 3.0 WIScrophulariaceae 4 1 0.6 1.1 28 SE 5.2 4 3.0 2.6 2 31 1 2.25 1.7 1.2 1.9 2.0 6.2Scrophulariaceae 5 0.8 1 SE 2 5 1.8 18.5 6Scrophulariaceae 6.0 0.9 BE 1.1 0.9 5.6 DicotScrophulariaceae 6.0 3 3 1.2 SE 1 0.4 28 1.3 5.1Scrophulariaceae 0.9 1 6.0 1.0 WI 1b Perennial 0.5 25.5 forb 1b 6.1Scrophulariaceae 0.0 Dicot 1.2 BE 1 5 24.5 0.0 0.7 2.3Scrophulariaceae Dicot 5 WI/IN 1Scrophulariaceae 1b 6.0 Perennial 1 1 SI forb 4.5 9Scrophulariaceae 1 4 5.0 1.1 BE/SI 0.9Scrophulariaceae 1 1 Perennial 27 Dicot forb SI 1.0 0.4 6.0 3.8 3.25 2.7 4 0.4 0.0 18.75 Dicot Perennial 13.5 3 6 4 forb 2.9 1 0.0 2.0 Dicot 5 1 14.25 Dicot 1.0 1b 4.5 0.5 5 Annual forb 0.9 0.3 4.0 1.4 1 Dicot 5 1 Annual 3.0 forb Annual 0.5 0.6 Dicot 2.1 forb 1 3.0 0.5 0.9 Annual forb 1.2 0.2 4 Annual forb 0.5 1 1 1 1 Dicot Dicot Dicot 1 Annual forb 1 Annual forb 1 1 1 Annual forb Dicot Dicot Dicot 1 Dicot 1 Annual forb Shrub Annual forb Dicot Annual forb Annual forb Scrophulariaceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 1 Dicot Perennial forb M. tenuis . elata ssp. ssp. var. neglecta spiralis ssp. ssp. ssp. ssp. ssp. ssp. (including ssp. 1 ssp. ONTINUED ) Taxon 1. C calyculatum pilosus brachiatus brevilobata lithospermoides rubicundula brunneus capillaris viscidus PPENDIX Antirrhinum subcordatum Antirrhinum vexillo- Antirrhinum virga Castilleja af®nis Castilleja foliolosa Castilleja hispida Castilleja miniata Castilleja minor Castilleja pruinosa Castilleja rubicundula Castilleja rubicundula Collinsia greenei Collinsia multicolor Collinsia sparsi¯ora Cordylanthus nidularius Cordylanthus pilosus Cordylanthus pringlei Cordylanthus tenuis Cordylanthus tenuis Cordylanthus tenuis Cordylanthus tenuis Keckiella lemmonii Mimulus douglasii Mimulus glaucescens Mimulus layneae A 2005] SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA 257 11 (rhiz.) (hemipar.) (hemipar.) (rhiz.) (rhiz.) Lifeform 10 Tax. Cat. Dicot Perennial forb Dicot Perennial forb 9 Geog. Dist. 11111 1111 KL NC BA SC SN 8 Rarity 7 SE 6 SD 5 Sources Med. 4 Sum 3 Mean 2 Family Aff Scrophulariaceae SEScrophulariaceae SI 6.0Scrophulariaceae SIScrophulariaceae 18 BEScrophulariaceae 2.5 BE/SIScrophulariaceae SIScrophulariaceae 2.7 7.5 3.7 5.0 WI 3Scrophulariaceae SESterculiaceae 8 11 30 3 6.0 2.8 2.3Verbenaceae BE/SI 0.0 11 6.1Violaceae 3.0 3 3 6 6.75Violaceae 0.0 2.0 BE 24.5 1.2 3.0 4.0 5.5 3 0.7 4 8 BE 4 0.6 4.8 0.6 1.3 SI 4 2.0 2.0 1 0.3 0.3 0.5 14.5 6.0 5.2 4 1.9 2.4 1 2.8 0.0 31 1.1 1.2 3 1.5 0.0 13.75 1b 1 1 1 2.4 4 4.0 6 1 5 1.2 1.2 1 1 1b 1 6.0 2.0 0.7 1 Dicot 1 1.3 1b 2.0 Dicot 1 0.5 0.9 1 1 Annual forb Dicot Dicot Perennial Dicot forb Dicot 1 Dicot Perennial forb Perennial 1 forb Dicot Perennial 1 forb Annual forb Dicot Perennial forb 1 Perennial forb Dicot Shrub Perennial Dicot forb Perennial forb Scrophulariaceae BE/SI 4.0 16 4 4.5 2.4 1.2 1 Dicot Perennial forb Scrophulariaceae SE 5.6 33.5 6 6.0 1.2 0.5 4 1 Dicot Annual forb ViolaceaeViolaceaeViolaceae BE/SIViolaceae WI 4.0Violaceae SI 16 2.3 BE 11.35 WI/IN 2.5 4 5.1 1.3 5 12.5 25.5 4.0 4 2.0 2.3 5 5 2.4 1.2 3.0 1.1 3 6.0 0.9 1.4 2.0 0.4 1 0.6 1.2 1 1 1b 0.7 1 1 1 1 1 Dicot Dicot Perennial forb Perennial Dicot forb 1 Dicot Perennial forb Perennial forb . azureus ssp. integrifolia var. ssp. lobata 1 ssp. ONTINUED ssp. Taxon 1. C decumbens ssp. linearifolius occidentalis PPENDIX Orthocarpus pachystachyus Pedicularis howellii Penstemon azureus Penstemon ®liformis Penstemon parvulus Penstemon purpusii Triphysaria ¯oribunda Veronica copelandii Fremontodendron californicum Verbena californica Viola cuneata Viola douglasii Mimulus nudatus Mimulus primuloides Viola hallii Viola lobata Viola ocellata Viola primulifolia Viola purpurea A