A DATABASE OF SERPENTINE AFFINITY › wildflowers › beauty › ...SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA: A DATABASE OF SERPENTINE AFFINITY H. D. SAFFORD1,2,J.H.VIERS3, AND

MADROÑO, 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. HARRISON21 USDA-Forest Service, Pacific Southwest Region, 1323 Club Drive,

Vallejo, CA [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 affinity to ultramafic (‘‘serpentine’’) sub-strates for taxa in the California flora, USA. We constructed our database through an extensive literaturesearch, expert opinion, field observations, and intensive use of accession records at key herbaria. Wedeveloped a semi-quantitative methodology for determining levels of serpentine affinity (strictly endemic,broadly endemic, strong ‘‘indicator’’, etc.) in the California flora. In this contribution, we provide a listof taxa having high affinity to ultramafic/serpentine substrates in California, and present information onrarity, geographic distribution, taxonomy, and lifeform. Of species endemic to California, 12.5% arerestricted to ultramafic substrates. Most of these taxa come from a half-dozen plant families, and fromonly one or two genera within each family. The North Coast and Klamath Ranges support more serpentineendemics than the rest of the State combined. 15% of all plant taxa listed as threatened or endangered inCalifornia show some degree of association with ultramafic substrates. Information in our database shouldprove valuable to efforts in ecology, floristics, biosystematics, conservation, and land management.

Key Words: serpentine, ultramafic, California, endemism, diversity.

INTRODUCTION

Ultramafic rocks, often called ‘‘serpentine’’ byecologists, botanists and pedologists, underlie morethan 6000 km2 of the land area of the State of Cal-ifornia (Harrison et al. 2000). The edges of conti-nental plates often include bands of these vestigesof oceanic mantle rock, accreted during the geolog-ic process of subduction, and later uplifted and ex-posed during mountain building and subsequenterosion. Ultramafic rocks and the soils that developon them are characterized by critically low levelsof most principal plant nutrients (N, P, K, Ca), andexceptionally high levels of Mg and Fe and a suiteof toxic trace elements including Cr, Ni, and Co.Outcrops of ultramafic rocks support high numbersof edaphic-endemic taxa throughout the world(Brooks 1987). The California serpentine flora isthe richest in the temperate zone, and consists ofhundreds of species and subspecies that are largelyor entirely confined to ultramafic substrates.

Serpentine endemism is a key feature of the di-versity of the California flora (Raven and Axelrod1978; Kruckeberg 2002). Of about 1410 full spe-cies endemic to the State (Hickman 1993), Kruck-eberg (1984) estimated that about 180 were endem-ic to serpentine. If these numbers are at least ap-proximately correct, then about 13% of the plantspecies endemic to California are serpentine en-demics. This is a remarkably high number whenone considers that only 1.5% of the State is under-

lain by ultramafic rocks (6000 km2/406,280 km2).In addition, because they tend to have small geo-graphic ranges and because many of them occur inthe rapidly urbanizing San Francisco Bay Area, ser-pentine endemics are overrepresented among thestate’s rare, sensitive, and listed plant taxa (Skinnerand Pavlik 1994). The ecology of California’s ser-pentine plants has been extensively studied at theUniversity of California’s Sedgwick Ranch Reserve(e.g., Seabloom et al. 2003; Gram et al. 2004) andMcLaughlin Reserve (e.g., Harrison et al. 2003;Safford and Harrison 2004) and Stanford Univer-sity’s Jasper Ridge Reserve (e.g., McNaughton1968; Huenneke et al. 1990; Hobbs and Mooney1991).

Botanists have relied for two decades on themonograph by Arthur Kruckeberg (1984) for mostof their information on Californian serpentine-en-demic plant taxa. Since then, publication of the Jep-son Manual (Hickman 1993), and a proliferation ofnew botanical research and name changes have leftthis list in need of updating. Our initial aim was tomodify Kruckeberg’s (1984) list, primarily usinginformation from Hickman (1993), to use in ourresearch on diversity patterns (Harrison et al. 2000,2004). However, it soon became clear that wewould have to expand and intensify our search forthe best available information. Complicating thiseffort, plants show a continuum in degrees of ser-pentine restriction, and are sometimes more restrict-

2005] 223SAFFORD ET AL.: SERPENTINE ENDEMISM IN THE CALIFORNIA FLORA

ed in some parts of their geographic ranges thanothers, thus contributing to inconsistencies amongreports from different sources. This led us to adopta semi-quantitative procedure for scoring plant taxaon their reported degree of serpentine affinity.

In this contribution, we present a summary of ourcurrent database of serpentine affinity in the Cali-fornia flora. The database was constructed via anextensive literature search, expert opinion, field ob-servations, web research, and intensive use of ac-cession records at key herbaria. It provides data onlevels of serpentine endemism, rarity, geographicdistribution, taxonomy, and lifeform.

METHODOLOGY

We began by conducting a database search of theelectronic Jepson Manual (Hickman 1993) main-tained by the Jepson Herbarium at the Universityof California-Berkeley (UC-JEPS 2004a). The da-tabase was queried for all taxa with ‘‘serpentine’’,‘‘ultramafic’’, or related (e.g., ‘‘asbestos soils’’) ref-erences in the habitat description. Taxa containing‘‘non-serpentine’’ in the description were removedafterward. We cross-checked the 391 serpentine-re-lated taxa found in the Jepson Manual with Kruck-eberg (1984), who listed those taxa he believed tobe endemic to ultramafic substrates in California,and those that were either local or regional ‘‘ser-pentine indicators’’ (i.e., nonendemic taxa whosedistributions are nonetheless skewed toward occur-rences on ultramafics). Taxonomic updates in theJepson Manual (Hickman 1993) were applied to theKruckeberg list (which included 377 taxa afterthese revisions), and then those taxa not on the Jep-son-derived list were added to our database. Thisresulted in a list of 529 taxa; of these, 287 were notshared between the two sources. We then added tothe list a number of taxa that we considered to belikely endemics or indicators but which were notindicated as such by either Kruckeberg (1984) orthe Jepson Manual (1993). Finally, published lit-erature (e.g., Meinke and Zika 1992; Nelson andNelson 2004; Baldwin 1999 and 2001; Barkley1999; Porter and Johnson 2000; Zika et al. 1998)and the online Jepson Interchange Jepson FloraProject (UC-JEPS 2004b) were consulted for tax-onomic revisions and taxa newly described sincethe publication of the Jepson Manual.

To score the affinity of taxa to ultramafic sub-strates, we adopted a modification of Kruckeberg’smeasures of ultramafic ‘‘fidelity’’. In his AppendixC, Kruckeberg (1984) used two or three ‘‘1’’s tosignify increasing levels of endemism: three ‘‘1’’swere attached to taxa with 95–100% of their oc-currences found on ultramafics, two ‘‘1’’s signifiedtaxa with 85–94% fidelity. In his Appendix D,Kruckeberg used one or two exclamation marks(‘‘!’’s) to signify increasing levels of fidelity to ul-tramafic substrates among supposed nonendemic‘‘indicator’’ taxa. In both appendices, question

marks (‘‘?’’) were attached to those taxa for whichmore information was necessary to confidently as-sign their status. Some of the ‘‘tentative’’ endemicswere included in the indicator appendix as well,thus these taxa occur twice in Kruckeberg’s lists.We combined Kruckeberg’s two scales, and addedtwo levels to yield six levels of ultramafic affinity,where 6 represents a ‘‘strict endemic’’ ($95% ofoccurrences on ultramafics), and successively lowervalues signify lower affinity to the substrate (5 585–94% of occurrences; 4 5 75–84%; 3 5 65–74%; 2 5 55–64%; 1 5 45–54%). By this defini-tion, ‘‘1’’ thus represents a species found about halfof the time on serpentine. We consider scores be-tween 1 and 2 to indicate ‘‘weak indicators’’, anda score of about 1 to mean an ‘‘indifferent’’ taxon.The Kruckeberg fidelity scale crosswalks to ours inthe following fashion: ‘‘111’’ 5 6; ‘‘11’’ 5 5;‘‘!!’’ 5 3; one ‘‘!’’ 5 2. Those taxa which occurredin both Kruckeberg’s endemic and indicator tableshad their two scores averaged: these all fell be-tween ‘‘3’’ and ‘‘4’’ on our scale. For example, Cu-pressus macnabiana was rated ‘‘111’’ in Kruck-eberg’s Appendix C (i.e., ‘‘6’’ on our scale), and‘‘!!’’ [i.e., ‘‘2’’ in our scale] in Appendix D; thesewere averaged to ‘‘4’’ on our scale.

We attached our categorical levels of ultramaficaffinity to all of the species in our hybrid Jepson-Kruckeberg database. In the case of the Kruckebergtaxa, we simply cross-walked the Kruckeberg fi-delity codes to our scale as described above, mak-ing some adjustments based on more recent taxo-nomic revisions and combinations. In the case ofthe Jepson Manual taxa, we were forced to interpretthe language used in habitat descriptions to deter-mine levels of affinity. We used the following in-terpretations of description language to assign af-finities: a ‘‘6’’ was assigned where the habitat de-scription categorically stated ‘‘serpentine’’ or ‘‘ul-tramafic’’ (a ‘‘5’’ if there was some indication thatthis restriction was not absolute); a ‘‘4’’ was as-signed where the modifiers ‘‘generally’’ or ‘‘usuallyserpentine’’ were used; ‘‘especially’’ or ‘‘often’’equaled ‘‘3’’; ‘‘sometimes’’ or ‘‘occasionally’’equaled ‘‘1’’. In a few cases, affinity levels wereassigned based on ancillary information in the hab-itat and/or range description rather than on explicitstatement of serpentine affinity.

We then conducted a broad survey of the litera-ture, regional botanical experts, and herbaria rec-ords to obtain as many sources as possible for eachtaxon in our database, and to add to the databaseany taxa we might have overlooked. We manuallyconsulted every species description in a variety ofregional and local floras (Clifton 2001; Ertter andBowerman 2004; Howell 1970; McMinn 1939; Os-wald 2002; Smith and Wheeler 1992), and guide-books to rare and sensitive taxa (Hanson 1999;Hoover et al. 1993; Jimerson et al. 1995; McCarten1988; McCarten and Rogers 1991; Nakamura andNelson 2001; Trinity SIPS 2001; USFWS 1998).

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We also consulted the CalFlora Online Species Da-tabase (CalFlora 2004), and the California NativePlant Society Online Inventory of Rare and Endan-gered Plants (CNPS 2004). We added columns toour database for each source, and gave scores (1–6, as described above) to each taxon for which ahabitat description suggested an ultramafic affinity.Information on serpentine affinity in the CalFloradatabase is limited to taxa from the Sierra Nevadaand to rare taxa statewide, and does not includesufficient information to determine degree of affin-ity (A. Dennis, personal communication). CalFlorawas therefore not treated as a typical ‘‘source’’, andCalFlora serpentine taxa were simply given a scoreof 0.5 to be added later to the sum of scores whenfinal ultramafic affinities were calculated (see be-low). The California Natural Diversity Database(CNDDB) was not searched, as we consulted all ofthe primary resources originally used to buildCNDDB, and the CNPS Online Inventory (seeabove) is updated from the same contemporarysources as CNDDB.

We calculated preliminary mean affinities fortaxa in our database by summing the scores acrosssource columns and adding the CalFlora score (ifpresent), then dividing by the number of sources(not including CalFlora) for the taxon in question.We also calculated the number of sources, the me-dian score, and the standard deviation and standarderror of the scores for each taxon. We then sent thedatabase to approximately 40 state and regional ex-perts for their review and input, and asked them toscore serpentine affinity using the 1–6 scale fortaxa with which they were familiar. These individ-uals included botanists employed by federal andstate land management agencies, universities, mu-seums, non-governmental organizations, and pri-vate consulting firms. We received 17 substantivereplies, and incorporated their input into an updateddatabase.

The next step was to ensure that we had at leastthree sources of serpentine affinity for each taxonin our database; given the great differences betweenthe Jepson Manual and Kruckeberg’s list, we felt athird opinion was important. We focused on thosetaxa for which we had less than three sources, aswell as those with high variability in scores. Webegan by consulting the habitat descriptions inMunz and Keck (1968) for every taxon in our da-tabase with less than three sources. We then turnedto Herbaria accession records. We searched the on-line ‘‘SMASCH’’ accession databases of the UCand Jepson Herbariums at UC-Berkeley at (UC-JEPS 2004c) for all taxa with one or two sources,and for all taxa with affinity-score standard devia-tions $ 1.0 (a total of 548 taxa). For any NorthernCalifornia taxa remaining with less than threesources and/or high variability, we then searchedthe online accession database of the Biological Sci-ences Herbarium at Chico State University (CSU-BSH 2004; a total of 164 taxa were searched).

In our online accession database research, wefollowed the following protocol:

1. We began with the most recent accession re-cords and worked backwards, as habitat descrip-tions before the mid 1970’s usually lack sufficientlydetailed information on substrate and location.

2. We consulted the habitat description for eachrecord. If the description included enough infor-mation to determine the substrate, we noted wheth-er it was ultramafic or non-ultramafic. We did notcount multiple accession records from the samecollecting trip and location as different records.

3. On the average, about ⅓ of the accession rec-ords consulted had sufficient information to deter-mine if a collection had been made on ultramaficsor not. Not all of these determinations were madesimply based on the collector’s habitat description.For example, many California counties do not con-tain outcrops of ultramafic rocks (e.g., Los Angeles,San Diego, San Bernardino, Modoc). Collectionsfrom these counties were coded as ‘‘nonserpentine’’even where habitat descriptions were missing. Also,collections from well-known collecting locations onultramafics (e.g., Blue Banks in Glenn County, RedButte in Siskiyou County, or the mouth of 18-MileCreek on the Middle Fork Smith River, Del NorteCounty) were coded as ‘‘serpentine’’ even wherehabitat descriptions were missing. Finally, wherewe had trouble getting a sufficient number of rec-ords with habitat descriptions, or where it was oth-erwise critical to get more information, we usedlocation information in the accession record (whereit existed) to do further research. We used TOPO!Software (National Geographic Maps 2000) to lo-cate coordinates or named locations and then con-sulted geological maps (ranging from 1:250,000 to1:25,000) to determine if the location was on anultramafic outcrop. Only those occurrences whichcould be confidently assigned to ultramafics wereidentified as such.

4. We continued until we had recorded habitatinformation from at least 10% of the total accessionrecords for the species in question. Our minimumwas 10 records, unless there were fewer than 10records with habitat descriptions and reasonably lo-catable site information (286/548 taxa had fewerthan 10). Our maximum was usually 20, althoughwe went beyond 20 in some cases.

5. We summarized the accession record resultsfor each taxon by dividing the total number of rec-ords with sufficient habitat or location informationto determine substrate by the number of recordsrecording serpentine/ultramafics, and then multi-plied the result by 100 to get a percentage. We thencross-walked the percent value to our scale of ul-tramafic affinity: 95–100% of records on ultramaf-ics 5 6; 85–94% 5 5; 75–84% 5 4; 65–74% 5 3;55–64% 5 2; 45–54% 5 1; 35–44% 5 0.75; 25–34% 5 0.5; 15–24% 5 0.25; .0–14% 5 0.1; 05 0.

Finally, T. Nelson and S. Carothers also used the


Humboldt State University Herbarium to provideinformation to us on a number of under-document-ed taxa from Northwestern California.

In our accession records research, we necessarilyassumed that: (1) the taxon itself was correctlyidentified on the accession record; (2) the substratewas correctly identified by the collector; and (3)ultramafic substrates were neither more nor lesslikely to be identified correctly (or at all) than othersubstrates. The last assumption is probably flawed,as serpentine and other ‘‘charismatic’’ substrates—given their close connection to plant endemic taxaand their relative ease of identification—are almostcertainly more likely to be identified than ‘‘normal’’substrates. This could theoretically lead to acces-sion records ‘‘overstating’’ the degree of a taxon’saffinity to ultramafic substrates. In practice, how-ever, we found that the accession records were gen-erally somewhat more conservative than our liter-ature sources vis-à-vis the serpentine affinities ofthe taxa in our database.

Our final database included 18 columns of infor-mation sources for serpentine affinity, plus a col-umn for CalFlora. We summed these affinity valuesand took their mean (not including CalFlora in thedenominator). We also calculated the mean withoutCalFlora, the median, the standard deviation, andthe standard error. We identified each taxon by tax-onomic category (pteridophyte, gymnosperm, dicotor monocot), and by lifeform (annual forb, peren-nial forb, annual graminoid, perennial graminoid,shrub, tree). For rare taxa, we added the rarity rat-ing from the California Native Plant Society OnlineDatabase of Rare and Endangered Plants (version6.04d, 11-12-2004). The following information wasalso added to the complete database: geographicdistribution in California for each taxon (by JepsonManual geographic subdivisions); elevational range(from Hickman 1993); the geographic distributionof, and number of species of the genus of eachtaxon (from Mabberly 1996); and the commonname (from Hickman 1993, and the Natural Re-source Conservation Service PLANTS online da-tabase [USDA-NRCS 2005]). Aside from a sum-mary of the geographic distribution, this informa-tion is not presented in the current paper, but isavailable on request from the first author, as are theaffinity values calculated for each source.

RESULTS

A summary table of the current database is pre-sented in Appendix 1. Appendix 1 includes 669taxa, ranging in affinity from 6.25 to 1.00 (somevalues exceed 6 because they were identified as ser-pentine taxa in the CalFlora Database). Our full da-tabase includes 698 taxa, 29 of which have meanserpentine affinities of , 1; we did not includethese taxa in the current paper. The greatest numberof sources we located for any single taxon was nine(four taxa). We found eight sources for eight taxa

and seven sources for 19 taxa; 587 taxa had be-tween three and six sources. Eighty-one taxa hadfewer than three sources (77 with two, three withone). Somewhat more than half of the taxa (387)in our original list had standard deviations for ser-pentine affinities . 1.0.

Since our serpentine affinities are calculated asthe means of multiple sources, our values fall on acontinuous scale, rather than in categories. Giventhis, we recognized taxa with mean affinities . 5.5as ‘‘strict endemics’’ (analogous to Kruckeberg’s‘‘111’’, or taxa with . 95% of their occurrenceson ultramafics), and taxa with mean affinities . 4.5and , 5.5 as ‘‘broad endemics’’ (analogous toKruckeberg’s ‘‘11’’, taxa with about 85–94% oftheir occurrences on ultramafics). Using these def-initions, 164 taxa are strict endemics, while 82 taxaare broad endemics, for a total of 246 endemic taxa;176 of these are full species. Among the remainingtaxa, 123 are ‘‘strong serpentine indicators’’(Kruckeberg 1984), with scores ranging from 2.5to 3.4 (about 65–74% of their occurrences on ul-tramafics); 150 are ‘‘weak indicators’’, falling be-tween 1.5 and 2.4 on our scale (6 55–64% of theiroccurrences on ultramafics); and 79 fall in a grayarea between weak indicators and indifferent taxa(between 1.0 and 1.4 on our scale, or about 50–54% of occurrences. Seventy-one taxa have affinityscores between 3.5 and 4.4 (about 75–84% of theiroccurrences on ultramafics), and thus represent thetransition from strong indicators to broad endemics.

Six families account for more than half of all theendemics: Asteraceae, Liliaceae, Brassicaceae, Po-lygonaceae, Scrophulariaceae, and Apiaceae (Table1). The 20 most important plant families among theserpentine endemics are shown in Fig. 1, with thepercentage of the serpentine endemic flora that theycontribute, as well as the percentage of the totalCalifornia endemic flora that they contribute. Fam-ilies that proportionally contribute more to the ser-pentine endemic flora than to the California endem-ic flora include Liliaceae, Brassicaceae, Polygona-ceae, Linaceae and Caryophyllaceae. Familieswhose level of endemism is much lower on serpen-tine than it is statewide include Fabaceae, Poaceae,Boraginaceae, and Rosaceae (Fig. 1).

The most diverse genera in our list of serpentineendemics are Streptanthus (Brassicaceae) and Er-iogonum (Polygonaceae), followed by Hesperoli-non (Linaceae) and Arctostaphylos (Ericaceae) (Ta-ble 2). There are 21 genera with at least four taxaamong the endemics. These represent 14 plant fam-ilies, with Asteraceae (four genera among the en-demics), Liliaceae (three genera), Scrophulariaceae(two genera) and Brassicaceae (two genera) theonly families with multiple genera in the list. Figure2 compares the contribution of these genera to theserpentine endemic flora with their contribution tothe California endemic flora. All but five or six ofthese genera have a greater level of endemism toserpentine than they have within the State as a


TABLE 1. NUMBERS OF SERPENTINE ENDEMIC AND NEARENDEMIC TAXA, BY FAMILY. 1 Strict endemics. 2 Strict en-demics plus broad endemics. 3 Strict and broad endemicsplus ‘‘near endemic’’ taxa (taxa transitional from strongindicators to broad endemics).

Family

Serpentine affinity score

$5.51 $4.52 $3.53Totaltaxa

Asteraceae 26 37 45 106Liliaceae 15 28 37 85Brassicaceae 21 26 31 46Polygonaceae 10 17 19 39Scrophulariaceae 9 14 18 37Apiaceae 7 10 13 32Linaceae 8 9 9 14Ericaceae 5 8 10 15Polemoniaceae 6 7 8 18Caryophyllaceae 5 7 8 18Fabaceae 4 7 10 24Lamiaceae 4 6 10 17Crassulaceae 5 5 7 13Rhamnaceae 4 5 6 14Campanulaceae 3 5 8 12Onagraceae 3 5 7 12Hydrophyllaceae 4 4 8 15Rubiaceae 3 4 4 8Convolvulaceae 1 4 5 6Cyperaceae 1 4 5 8Poaceae 1 3 3 19Portulacaceae 0 3 5 16Boraginaceae 2 2 3 10Gentianaceae 2 2 2 3Iridaceae 2 2 2 4Malvaceae 2 2 2 5Salicaceae 2 2 2 3Garryaceae 1 2 2 2Rosaceae 1 2 5 10Cupressaceae 0 2 3 6Violaceae 0 2 3 7Asclepiadaceae 1 1 1 1Berberidaceae 1 1 1 4Dryopteridaceae 1 1 1 2Fagaceae 1 1 1 3Lentibulariaceae 1 1 1 1Papaveraceae 1 1 1 5Ranunculaceae 1 1 3 6Orchidaceae 0 1 1 3Pteridaceae 0 1 1 4Verbenaceae 0 1 1 1Cistaceae 0 0 0 1Orobanchaceae 0 0 0 1Pinaceae 0 0 1 6Plantaginaceae 0 0 0 1Polygalaceae 0 0 0 1Primulaceae 0 0 0 1Sarraceniaceae 0 0 1 1Saxifragaceae 0 0 1 2Sterculiaceae 0 0 0 1

Totals 164 246 315 669

whole. These genera include Streptanthus, Hesper-olinon, Lomatium and Minuartia. Only one genus(Phacelia) contributes less to the serpentine endem-ic flora than it does to the State as a whole; Arc-

tostaphylos contributes a similar percentage to bothfloras (Fig. 2).

Of the taxa in our database, there are 532 dicots(of which 204 are endemic), 119 monocots (38 en-demics), 12 gymnosperms (2 endemics) and sixpteridophytes (2 endemics). 207 taxa are annualforbs (of which 71 are endemics, including 7 of 14that can also be perennial/biennial), 383 are peren-nial forbs (150 endemics, including the 7 ‘‘annu-als’’ and 6 taxa which can also be shrubs), 24 areperennial graminoids (7 endemics), 64 are shrubs(23 endemics, including 6 taxa shared with the pe-rennial forbs and 1 which assumes both tree andshrub forms), and 12 are trees (2 endemics) (Ap-pendix 1). Of the endemic perennial forbs, 24 arebulb plants (all Liliaceae), 17 are rhizomatous(from ten different Families), three are hemipara-sites (Scrophulariaceae), and one is carnivorous(Lentibulariaceae) (Appendix 1).

Using Kruckeberg’s (1984) physiographic prov-inces of California (which correspond more or lessto major geographic subdivisions mapped in theJepson Manual (Hickman 1993)), we found the fol-lowing geographic distribution of serpentine en-demic taxa (Fig. 3): The North Coast, consideredin toto (i.e., the Jepson Manual’s NCo and NCoRsubregions (Hickman 1993)), supports approxi-mately 118 serpentine endemics, with 49 of theserestricted to that area. The Klamath Region (JepsonManual subregion KR), supports 98 endemic taxa,with 54 restricted to that area (including taxa alsofound in neighboring SW Oregon). The San Fran-cisco Bay Area (Jepson Manual subregion SnFrBplus the sections of NCo and CCo bordering it)supports about 51 endemics, with 24 found onlythere. The South Coast Ranges, including the Chan-nel Islands and the Santa Ana Mountains (i.e., Jep-son Manual subregions CCo, SCoR plus the fewultramafic outcrops that occur in the Jepson SWRegion), support 43 total endemics with 24 restrict-ed to that area. The Sierra Nevada (Jepson Manualregion SN) support 38 total serpentine endemictaxa, with 21 taxa restricted to the Range (Fig. 3).

Of the 669 taxa in our database, 295 are listedas ‘‘rare’’ or ‘‘uncommon’’ by the California NativePlant Society (CNPS) (Appendix 1). These include194 of the 246 taxa that we consider to be eitherstrict or broad serpentine endemics. One serpentineendemic taxon, Arctostaphylos hookeri subsp. fran-ciscana, is extinct in the wild and survives only incultivation. Of the 295 rare or uncommon taxa, 154are on CNPS List 1b, which lists plants consideredthreatened or endangered by either the State or Fed-eral governments, as well as unlisted plants whichCNPS considers rare enough to warrant listing; 111of these List 1b plants are serpentine endemics byour definition. Nine taxa (seven endemics) fromAppendix 1 are on CNPS list 2, which containsplant taxa that are rare in California but are notrestricted completely to the State; all of these taxaare either State listed and threatened or endangered,


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 flora in California,and to the California endemic flora as a whole.

TABLE 2. GENERA WITH MORE THAN THREE TAXA ENDEMICTO SERPENTINE.

Genus FamilyEndemic

taxa

Streptanthus Brassicaceae 18Eriogonum Polygonaceae 14Hesperolinon Linaceae 9Arctostaphylos Ericaceae 8Allium Liliaceae 7Lomatium Apiaceae 7Packera (Senecio) Asteraceae 6Calochortus Liliaceae 5Cordylanthus Scrophulariaceae 5Arabis Brassicaceae 4Calystegia Convolvulaceae 4Carex Cyperaceae 4Castilleja Scrophulariaceae 4Cirsium Asteraceae 4Erigeron Asteraceae 4Fritillaria Liliaceae 4Galium Rubiaceae 4Lessingia Asteraceae 4Minuartia Caryophyllaceae 4Monardella Lamiaceae 4Phacelia Hydrophyllaceae 4

or are eligible for listing. Eight taxa (four endem-ics) in Appendix 1 are found on CNPS list 3, whichlists uncommon taxa for which more information isrequired. Of taxa in Appendix 1, 123 (71 endemics)are on CNPS list 4, which contains taxa of ‘‘limiteddistribution or infrequent throughout a broader areain California’’.

DISCUSSION

In 1984, Kruckeberg estimated that the serpen-tine endemic flora of California numbered approx-imately 220 taxa (about 180 full species), and thata further 230 taxa were sufficiently associated withultramafics to be ‘‘indicators’’ of the substrate.Thus, Kruckeberg believed that about 450 taxawere associated with serpentine in California. Al-though our results suggest that the number of ser-pentine-associated taxa is closer to 670, they alsosuggest that Kruckeberg’s (1984) estimate of thenumber of full-species endemics was remarkablyaccurate (180 vs. 176). As Kruckeberg’s numbersalso suggested, serpentine endemics therefore com-prise approximately 12.5% (176/1410) of the plantspecies endemic to California. Based on numbersfrom the Jepson Manual (Hickman 1993; R. Moepersonal communication), the percentage of serpen-tine endemics among California endemic species,subspecies and varieties is about 11.4% (246/2153).

Kruckeberg’s (1984) estimates of endemics byCalifornia geographic region are somewhat less ac-curate than his statewide estimate (see Fig. 3), butKruckeberg’s data sources in the 1970’s and early1980’s were extremely limited compared to ours.As did Kruckeberg, we found that the North CoastRanges support more serpentine endemics plantsthan any other geographic region, but that theKlamath Ranges (and adjoining SW Oregon) sup-port many more restricted endemics than Krucke-berg thought was the case (54 vs. 30). Kruckeberg’sestimates for the numbers of restricted endemics in


FIG. 2. The twenty-one most important genera of serpentine endemic plants (i.e., including strict and broad serpentineendemics), with the percentage of endemic species they contribute to the serpentine endemic flora in California, andto the California endemic flora as a whole.

FIG. 3. Geographic distribution of serpentine endemictaxa in California. ‘‘Total endemics’’ includes all Califor-nia serpentine endemic taxa present in a given region;‘‘restricted endemics’’ includes only those taxa restrictedto a given region; black bars represent Kruckeberg’s(1984) estimates of restricted endemics.

the North Coast Ranges and the Bay Area are verysimilar to our numbers (Fig. 3), but he overesti-mated the number of endemics in the South CoastRanges (36 vs. 24). Kruckeberg estimated that ei-ther 13 or 16 (depending on whether one goes bythe text or the tables in Appendix E) endemic taxawere restricted to the Sierra Nevada; we found 21taxa thus restricted.

Reasons for differences between our numbersand Kruckeberg’s (1984) are many, but belong totwo broad categories. The primary reason is qualityand quantity of information. In many cases, Kruck-eberg’s information had to come through his ownfield experience, or through hard copy herbariumrecords, which—before the late 1970’s—were no-

toriously uninformative when it came to habitat de-scription. In contrast, many data sources we ac-cessed were available electronically and could bequeried and retrieved remotely.

The other principal reason for difference is theinevitable discoveries and taxonomic reorganiza-tions that occur over a 20-year period. Krucke-berg’s work came before publication of the JepsonManual (Hickman 1993), which contained manysignificant changes in California plant taxonomy. Aconsiderable number of serpentine endemic taxa inthe Jepson Manual were wholly unknown to Kruck-eberg in 1984. Examples include Calochortusraichei S. Farwig & V. Girard, Minuartia stoloni-fera T. W. Nelson & J. P. Nelson, Perideridia ba-cigalupii Chuang & Constance, and Monardellastebbinsii Hardham & J. Bartel. Since the Manual’spublication, there have been further changes (e.g.,Barkley 1999; Baldwin 1999; Porter and Johnson2000). Serpentine endemic taxa named since pub-lication of the Jepson Manual include Harmoniaguggolziorum B. G. Baldwin, Carex serpentinicolaP. F. Zika, and Silene serpentinicola T. W. Nelson& J. P. Nelson.

As a null hypothesis, one might expect that thedistribution of endemic plant taxa across plant fam-ilies and genera on California serpentines wouldmore or less mirror the distribution of endemics inthe State as a whole. Our data demonstrate that thisassumption is incorrect at both taxonomic levels,but the root of this difference seems to be largelyat the level of genus. A number of families con-tribute a much higher proportion of the serpentine


TABLE 3. EXAMPLES OF ‘‘REGIONAL’’ SERPENTINE INDICATORS, SENSU KRUCKEBERG (1984). 1 CA 5 California, KL 5Klamath Ranges, NC 5 North Coast Ranges, BA 5 San Francisco Bay Area, SC 5 South Coast Ranges, SN 5 SierraNevada.

Taxon Distribution1 Comments1

Allium amplectens CA SC—broad endemic; Northern CA—weak indicator at bestAspidotis densa CA Marin County—broad endemic; rest of NC—weak to strong indicator;

KL—broad endemic to strong indicator, depending on locality; restof CA—strong indicator

Festuca californica CA Northern SN and KL—strong indicator to broad endemic; NC—primar-ily weak indicator; rest of CA—indifferent

Lupinus onustus KL, SN KL—broad endemic; SN—indifferent (mostly non-ultramafic)Pinus attenuata CA Mendocino County and neighboring NC—broad endemic; rest of NC

and SC—strong indicator; KL—weak indicator; SN—weak indicatorto indifferent

Pinus jeffreyi KL, NC, SC, SN KL and NC—6 strict endemic; Westslope of northern SN—strong indi-cator; rest of CA—indifferent

Quercus vaccinifolia KL, NC, SN Mendocino County and neighboring NC—broad endemic; Northern NCand KL—weak indicator; SN—indifferent

Sedum obtusatumssp. obtusatum

KL, SN KL and NC—6 broad endemic; SN—weak indicator or indifferent

Stachys pycnantha CA Marin County—broad endemic to strong indicator; Northern SN—veryweak indicator; rest of CA—weak indicator or indifferent

Viola douglasii CA Plumas County—endemic; NC—strong indicator; rest of CA—indifferent

endemic flora than they do of the California endem-ic flora (Fig. 1), but our database shows that mostof these ‘‘anomalies’’ are due to one or two generawithin those families (see Fig. 2). Examples includeFritillaria and Allium in Liliaceae, Minuartia inCaryophyllaceae, Streptanthus and Arabis in Bras-sicaceae, Hesperolinon in Linaceae, and Eriogon-um in Polygonaceae. Many of these genera arewell-known as foci of neoendemism (i.e., generawith groups of actively and rapidly speciating taxa)(Raven and Axelrod 1978). It is interesting thatsuch prominent California plant families like Scro-phulariaceae, Hydrophyllaceae, Boraginaceae, On-agraceae and Polemoniaceae are underrepresentedon serpentine substrates. Certain highly diversegenera in California are also proportionally under-represented as serpentine endemics (e.g., Clarkia,Phacelia, Ceanothus, Gilia, and Mimulus).

As we constructed our database, taxa with highvariability in serpentine affinity scores were taggedfor further research (e.g., through accession rec-ords; see Methodology) so that we might be ableto discern taxa that truly varied geographically intheir affinities from taxa that simply suffered frominadequate or faulty information. The former werecalled ‘‘regional indicators’’ by Kruckeberg (1984),i.e., taxa that are considered serpentine endemics orindicators in one part of their range but show lessor no affinity for ultramafic substrates in other partsof their range. In his Appendix D, Kruckeberg(1984) tried to summarize where the different re-gional indicators he had identified occurred on ul-tramafics. We refer the reader to Kruckeberg (1984)for details on these taxa (most of which also occurin our database), but most regional indicators in ourdatabase can be recognized by searching for taxawith: (1) relatively wide geographic distributions,

(2) lower mean serpentine affinity scores, and (3)high standard deviations in their affinity scores. Ta-ble 3 lists ten examples of regional indicator taxain our database.

Some of the variability in our serpentine affinityscores is thus due to geographic variation in affin-ities, but some is also due to inadequate, statisti-cally biased, or even faulty information from oursources. We attempted to offset these sources ofvariability by including as many sources as possiblein our database (and by using accession records),but were not successful in all cases. We considerany taxon with a standard deviation in affinity score. 1.5, or having fewer than three sources, as beingin ‘‘need of further research’’; this includes abouta third of the taxa in our database. Examples ofsuch taxa include: Lupinus lapidicola—called astrict serpentine endemic by Kruckeberg (1984) anda strong serpentine indicator by CNPS (2004), andwith 2/2 accession records in SMASCH with ultra-mafic habitat descriptions, but stated as occurringonly on granites by the Jepson Manual (Hickman1993) and Munz and Keck (1973); Phacelia pha-celioides—Kruckeberg (1984) and V. Yadon (per-sonal communication) believe this is a strict endem-ic, but the Jepson Manual is mute on the subject,and only 1/3 accession records in SMASCH are onultramafics (but the two nonserpentine locationsmay have misidentified geology given the location);and Allium lacunosum var. lacunosum—both theJepson Manual and Kruckeberg rate this as a strictendemic, Munz and Keck score it a strong indica-tor, but SMASCH has only 1/6 records on ultra-mafics.

Some species had surprising levels of ultramaficaffinity. For example, our database includes a num-ber of taxa that we personally have only rarely seen


on serpentine (e.g., Lathyrus vestitus var. vestitus,Apiastrum angustifolium, Emmenanthe pendulifloravar. penduliflora). It also includes other taxa whichwe would have characterized as being clearly in-different to ultramafic substrates, but which scoredhigher based on our sources (e.g., Adenostoma fas-ciculatum, Pinus balfouriana ssp. balfouriana). Asnoted above, some of these ‘‘discrepancies’’ maybe due to inadequate or biased data—the ultramaficaffinity of these types of taxa will drop as we col-lect more information. Many of these surprising af-finities are probably real however, and they are sim-ply a sign of our limited knowledge of the relation-ships between California plant life and ultramaficsubstrates.

In accession records, and in the literature, bota-nists and ecologists frequently misidentified basicrock types. For example, in accession records wefound a number of examples of peridotite beingcalled ‘‘volcanics’’ or even ‘‘sandstones’’. We alsofound multiple examples, in accession records aswell as in the literature, of gabbro and other basicintrusive rocks being misidentified as ultramafics.Gabbro and ‘‘basic’’ rocks are ‘‘mafic’’ in compo-sition—that is to say, they usually contain visiblefeldspars and they are geochemically distinct fromultramafic rocks. For example, the average alkali-gabbro contains 4–5 times as much Na as perido-tite, 5–10 times as much P, 3–4 times as much Kand Ca, and about ⅓ as much Mg (Ehlers and Blatt1982). The famous gabbro outcrops of EldoradoCounty (Pine Hill) or San Diego County are there-fore not ultramafic, even though the effect of thesubstrate on plant physiognomy and communitycomposition may appear similar. A number of spe-cies in our database appear to be primarily, if notexclusively gabbro endemics, but we lacked suffi-cient information to remove them from our list.These include Acanthomintha ilicifolia, Fremonto-dendron californicum ssp. decumbens, and Calo-chortus weedii var. vestus.

As has been frequently noted (Mason 1946a, b;Raven and Axelrod 1978; Kruckeberg 1984, 2002;Skinner and Pavlik 1994; McCarten 1997), Cali-fornia’s ultramafic soils support a very high pro-portion of the State’s rare plants. Based on our da-tabase, almost 11% (111/1021) of California’s rareplant taxa are either broadly or strictly restricted toultramafic substrates; 15% of List 1b taxa (154/1021) show high affinity for ultramafic substrates(i.e., they are endemics or indicators). In north-western California, 15% of plant taxa managed as‘‘sensitive’’ by the Forest Service are serpentine en-demics, and fully 30% are closely associated withultramafics (J. K. Nelson and L. Hoover personalcommunication). In 2002, Kruckeberg wrote that‘‘preservation of serpentine habitats in California isspotty, inadequate, and largely coincidental’’. Giv-en the great importance of ultramafic substrates tothe richness and distinctiveness of the Californiaflora, the conservation of these unique habitats

should be a high priority for land managementagencies and private conservation organizationsthroughout the State.

Our database of serpentine affinity updates, andexpands on the widely-used tables of serpentine en-demic and ‘‘indicator’’ taxa published in 1984 byArt Kruckeberg in his classic monograph on Cali-fornia serpentine ecology. Our data are also a quan-titative synthesis of the qualitative (and usually in-complete) allusions to serpentine affinity containedin habitat descriptions in California floras and floradatabases, including Munz and Keck (1973), Hick-man (1993), Oswald (2002), the online CalFloraDatabase (CalFlora 2004), and the California Na-tive Plant Society Online Inventory of Rare andEndangered Plants (CNPS 2004). Our data on ser-pentine endemism should prove valuable to effortsin ecology, biosystematics (Baldwin 1995), conser-vation, and land management. In particular, wehope that our database will help us better under-stand the nature and degree of serpentine endemismin the California flora, and we hope it will spur thecollection of additional, critical information neces-sary for conserving the rare plants and habitats ofultramafic substrates.

ACKNOWLEDGEMENTS

We would like to thank the following individuals forproviding information on plant affinities to ultramafic sub-strates: Joe Callizo, Sydney Carothers, Susan Erwin, Lin-nea Hanson, Lisa Hoover, David Isle, Marla Knight, NiallMcCarten, John McRae, Julie Kierstad Nelson, Tom Nel-son, Robert Preston, Barbara Williams, and Vern Yadon.Dick Moe of the Jepson Herbarium provided us with dataon identities and numbers of taxa endemic to California.Special thanks to Art Kruckeberg and Niall McCarten forcomments on early versions of the manuscript.

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APPENDIX 1. PLANT TAXA WITH HIGH AFFINITY TO ULTRAMAFIC SUBSTRATES IN CALIFORNIA. Ordered by family and taxon. 1 Names as in Hickman (1993); names in parenthesesare based on more recent revisions (see text for sources). 2 Affinity: SE 5 strict endemic, BE 5 broad endemic, BE/SI 5 broad endemic/strong indicator, SI 5 strongindicator, WI 5 weak indicator, WI/IN 5 weak indicator/indifferent. 3 Mean affinity score, including information from CalFlora. 4 Sum of all affinity scores, includingCalFlora. 5 Median of affinity scores. 6 Standard deviation of affinity scores. 7 Standard error of affinity scores. 8 California Native Plant Society rarity codes, from CNPSInventory of Rare and Endangered Plants of California, 11-2004. 9 Geographic distribution: KL 5 Klamath Ranges, NC 5 North Coast Ranges, BA 5 San Francisco BayArea, SC 5 South Coast Ranges, SN 5 Sierra Nevada. 10 Taxonomic category. 11 carn 5 carnivorous, cesp 5 cespitose, hemipar 5 hemiparasitic, paras 5 parasitic, rhiz 5rhizomatous. See text for more information.

Taxon1 Family Aff2 Mean3 Sum4 Sources Med.5 SD6 SE7 Rarity8

Geog. Dist.9

KL NC BA SC SNTax.Cat.10 Lifeform11

Angelica tomentosa Apiaceae SI 2.7 8 3 3.0 1.5 0.9 1 1 1 Dicot Perennial forbApiastrum angustifolium Apiaceae WI 1.5 7.6 5 0.1 2.5 1.1 1 1 1 1 Dicot Annual forbLigusticum californicum Apiaceae WI/IN 1.4 5.75 4 1.4 1.3 0.7 1 1 1 Dicot Perennial forbLomatium ciliolatum Apiaceae SE 6.0 18 3 6.0 0.0 0.0 1 1 1 Dicot Perennial forbLomatium congdonii Apiaceae SE 6.2 18.5 3 6.0 0.0 0.0 1b 1 Dicot Perennial forbLomatium dasycarpum ssp.

dasycarpumApiaceae BE/SI 3.6 21.5 6 3.5 1.9 0.8 1 1 1 Dicot Perennial forb

Lomatium engelmannii Apiaceae SE 5.8 34.5 6 6.0 0.8 0.3 4 1 1 Dicot Perennial forbLomatium hooveri Apiaceae SE 5.9 29.5 5 6.0 0.4 0.2 4 1 Dicot Perennial forbLomatium howellii Apiaceae SE 6.1 24.5 4 6.0 0.0 0.0 4 1 Dicot Perennial forbLomatium macrocarpum Apiaceae SI 2.7 8 3 3.0 0.6 0.3 1 1 1 1 1 Dicot Perennial forbLomatium marginatum Apiaceae BE 5.0 25 5 6.0 1.4 0.6 1 1 1 Dicot Perennial forbLomatium observatorium Apiaceae WI/IN 1.4 2.75 2 1.4 0.9 0.6 1 1 Dicot Perennial forbLomatium parvifolium Apiaceae SI 3.3 13 4 3.0 2.3 1.1 4 1 Dicot Perennial forbLomatium repostum Apiaceae SI 3.2 12.6 4 3.0 2.4 1.2 4 1 Dicot Perennial forbLomatium tracyi Apiaceae SE 6.1 42.5 7 6.0 0.0 0.0 4 1 1 Dicot Perennial forbLomatium triternatum var.

triternatumApiaceae SI 2.8 11 4 2.0 2.4 1.2 1 1 Dicot Perennial forb

Lomatium utriculatum Apiaceae WI 1.7 8.5 5 1.0 1.4 0.6 1 1 1 1 1 Dicot Perennial forbPerideridia bacigalupii Apiaceae BE 4.6 23 5 6.0 2.4 1.1 4 1 Dicot Perennial forbPerideridia kelloggii Apiaceae WI 2.1 10.6 5 2.0 2.0 0.9 1 1 1 1 Dicot Perennial forbPerideridia leptocarpa Apiaceae SE 5.6 22.5 4 6.0 1.0 0.5 4 1 Dicot Perennial forbPerideridia oregana Apiaceae WI 1.7 5 3 1.0 1.2 0.7 1 1 1 1 Dicot Perennial forbPerideridia pringlei Apiaceae BE/SI 3.7 18.5 5 3.0 2.3 1.0 4 1 Dicot Perennial forbSanicula bipinnatifida Apiaceae WI 1.8 7.1 4 2.0 1.5 0.7 1 1 1 1 1 Dicot Perennial forbSanicula hoffmannii Apiaceae WI 1.8 3.5 2 1.5 2.1 1.5 4 1 Dicot Perennial forbSanicula maritima Apiaceae WI 2.3 4.5 2 2.0 0.0 0.0 1b 1 1 Dicot Perennial forbSanicula peckiana Apiaceae BE 5.3 26.5 5 6.0 1.3 0.6 4 1 Dicot Perennial forbSanicula tracyi Apiaceae WI 2.1 8.5 4 1.0 2.6 1.3 1 Dicot Perennial forbSanicula tuberosa Apiaceae WI/IN 1.3 3.75 3 1.0 0.7 0.4 1 1 1 Dicot Perennial forbTauschia glauca Apiaceae BE/SI 3.5 10.5 3 3.0 0.6 0.3 4 1 Dicot Perennial forbTauschia hartwegii Apiaceae WI/IN 1.3 4 3 1.0 1.5 0.9 1 1 1 Dicot Perennial forbTauschia howellii Apiaceae WI 2.3 7 3 1.0 3.2 1.9 1 Dicot Perennial forbTauschia kelloggii Apiaceae SI 2.6 12.75 5 2.0 2.2 1.0 1 1 1 Dicot Perennial forb

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Asclepias solanoana Asclepiadaceae SE 6.0 42 7 6.0 0.0 0.0 4 1 Dicot Perennial forbAgoseris heterophylla Asteraceae WI/IN 1.4 4.1 3 1.0 1.5 0.9 1 1 1 1 1 Dicot Annual forbAncistrocarphus filagineus Asteraceae SI 3.3 13 4 3.0 0.5 0.3 1 1 1 1 1 Dicot Annual forbAntennaria argentea Asteraceae WI 1.9 7.75 4 0.8 2.7 1.4 1 1 Dicot Perennial forbAntennaria suffrutescens Asteraceae SE 5.6 22.5 4 6.0 1.0 0.5 4 1 1 Dicot Perennial forbArnica cernua Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 4 1 Dicot Perennial forb

(rhiz.)Arnica spathulata Asteraceae SE 5.5 16.5 3 6.0 1.2 0.7 4 1 Dicot Perennial forb

(rhiz.)Aster oregonensis Asteraceae WI/IN 1.1 3.25 3 1.0 0.9 0.5 1 1 1 Dicot Perennial forbBalsamorhiza macrolepis var.

macrolepisAsteraceae SI 2.5 15 6 2.0 1.1 0.5 1b 1 1 Dicot Perennial forb

Balsamorhiza sericea Asteraceae SE 6.2 18.5 3 6.0 0.0 0.0 1b 1 Dicot Perennial forbBrickellia greenei Asteraceae BE/SI 3.7 11 3 4.0 0.6 0.3 1 1 1 Dicot Perennial forbCacaliopsis nardosmia Asteraceae WI/IN 1.3 4 3 2.0 1.2 0.7 1 1 Dicot Perennial forbCalycadenia multiglandulosa Asteraceae SI 3.1 15.5 5 3.0 1.2 0.6 1 1 1 1 Dicot Annual forbCalycadenia oppositifolia Asteraceae SI 2.6 18 7 2.0 1.6 0.6 1b 1 Dicot Annual forbCalycadenia pauciflora Asteraceae BE 5.3 21 4 5.5 1.0 0.5 1 Dicot Annual forbCalycadenia truncata Asteraceae WI 2.1 12.5 6 2.5 1.1 0.5 1 1 1 1 1 Dicot Annual forbChaenactis glabriuscula var.

glabriusculaAsteraceae WI 1.7 5.1 3 2.0 1.5 0.9 1 1 1 Dicot Perennial forb

Chaenactis glabriuscula var.heterocarpha

Asteraceae SI 2.5 10 4 2.5 0.6 0.3 1 1 1 1 Dicot Annual forb

Chaenactis suffrutescens Asteraceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 Dicot Perennial forbChrysothamnus nauseosus ssp.consimilis

Asteraceae WI 1.8 8.85 5 2.0 1.5 0.7 1 1 1 Dicot Shrub

Cirsium andrewsii Asteraceae WI 1.7 5 3 2.0 0.9 0.5 1b 1 Dicot Perennial forbCirsium cymosum Asteraceae SI 3.0 12 4 2.0 2.0 1.0 1 1 1 1 1 Dicot Perennial forbCirsium douglasii var. breweri Asteraceae SI 3.0 12 4 3.0 1.6 0.8 1 1 1 Dicot Perennial forbCirsium fontinale var. campylon Asteraceae SE 5.9 29.5 5 6.0 0.4 0.2 1b 1 1 Dicot Perennial forbCirsium fontinale var. fontinale Asteraceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 1 Dicot Perennial forbCirsium fontinale var.

obispoenseAsteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Perennial forb

Cirsium hydrophilum var.vaseyi

Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 1 Dicot Perennial forb

Cirsium remotifolium Asteraceae WI/IN 1.0 3.1 3 1.0 1.0 0.5 1 1 1 Dicot Perennial forbCoreopsis stillmanii Asteraceae SI 2.7 8 3 3.0 0.6 0.3 1 1 1 Dicot Annual forbCrepis pleurocarpa Asteraceae WI 2.0 10 5 2.0 0.7 0.3 1 1 1 Dicot Perennial forbEricameria arborescens Asteraceae WI/IN 1.3 4 3 1.0 1.5 0.9 1 1 1 1 Dicot ShrubEricameria greenei Asteraceae WI 2.0 8.1 4 1.5 2.1 1.1 1 1 1 Dicot ShrubEricameria ophitidis Asteraceae SE 5.5 38.5 7 6.0 1.0 0.4 4 1 1 Dicot Shrub

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Erigeron angustatus Asteraceae SE 5.7 28.5 5 6.0 0.9 0.4 1b 1 Dicot Perennial forbErigeron bloomeri var. nudatus Asteraceae SE 6.2 18.5 3 6.0 0.0 0.0 2 1 Dicot Perennial forbErigeron cervinus Asteraceae SI 3.3 10 3 4.0 3.1 1.8 1 Dicot Perennial forb

(rhiz.)Erigeron decumbens var.

robustiorAsteraceae WI 1.5 4.5 3 2.0 1.2 0.7 4 1 Dicot Perennial forb

Erigeron foliosus var. confinis Asteraceae BE/SI 3.7 11 3 3.0 1.2 0.7 1 Dicot Perennial forbErigeron lassenianus var.

deficiensAsteraceae WI 1.7 5 3 2.0 1.5 0.9 1 Dicot Perennial forb

Erigeron petrophilus var.sierrensis

Asteraceae BE 4.8 28.5 6 6.0 2.1 0.8 4 1 Dicot Perennial forb(rhiz.)

Erigeron petrophilus var.viscidulus

Asteraceae WI 2.4 9.5 4 2.0 0.5 0.3 4 1 Dicot Perennial forb(rhiz.)

Erigeron reductus Asteraceae WI 2.0 8 4 2.0 1.6 0.8 1 1 1 Dicot Perennial forbErigeron serpentinus Asteraceae SE 6.2 18.5 3 6.0 0.0 0.0 1b 1 Dicot Perennial forbEriophyllum confertiflorum var.

tanacetiflorumAsteraceae WI 1.9 3.75 2 1.9 1.6 1.1 1 Dicot Shrub

Eriophyllum jepsonii Asteraceae BE/SI 3.5 17.5 5 3.0 1.5 0.7 4 1 1 Dicot ShrubEriophyllum lanatum var.

achillaeoidesAsteraceae WI 2.3 7 3 2.0 0.6 0.3 1 1 1 1 1 Dicot Shrub

Eriophyllum lanatum var.lanceolatum

Asteraceae WI 1.7 5 3 2.0 1.5 0.9 1 1 Dicot Shrub

Eriophyllum latilobum Asteraceae SE 5.5 16.5 3 6.0 1.2 0.7 1b 1 1 Dicot ShrubGrindelia hirsutula var. davyi Asteraceae WI 1.8 5.25 3 2.0 1.4 0.8 1 1 Dicot Perennial forbGrindelia hirsutula var.

hirsutulaAsteraceae WI/IN 1.2 3.6 3 1.0 1.0 0.5 1 1 Dicot Perennial forb

Grindelia hirsutula var.maritima

Asteraceae WI 1.7 5 3 2.0 0.9 0.5 1b 1 Dicot Perennial forb

Gutierrezia californica Asteraceae WI 1.8 5.25 3 2.0 1.4 0.8 1 1 1 Dicot Perennialforb, Shrub

Harmonia guggolziorum Asteraceae SE 6.0 18 3 6.0 0.0 0.0 1b 1 Dicot Annual forbHazardia stenolepis Asteraceae WI 2.0 6.1 3 3.0 1.7 1.0 1 Dicot Shrub (stem

succulent)Hazardia whitneyi var.

discoideaAsteraceae WI 1.9 5.75 3 2.0 1.1 0.7 1 Dicot Perennial

forb, ShrubHazardia whitneyi var. whitneyi Asteraceae WI/IN 1.0 2 2 1.0 1.4 1.0 1 Dicot Perennial

forb, ShrubHelenium bigelovii Asteraceae SI 2.9 11.5 4 2.5 2.3 1.2 1 1 1 1 Dicot Perennial forbHelianthus exilis Asteraceae SE 5.7 45.5 8 6.0 1.1 0.4 4 1 1 1 1 Dicot Annual,

Perennialforb

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Hemizonia congesta ssp.calyculata

Asteraceae WI 1.5 4.5 3 2.0 1.2 0.7 4 1 Dicot Annual forb

Hemizonia congesta ssp.congesta

Asteraceae WI/IN 1.3 4 3 2.0 1.2 0.7 1 1 1 Dicot Annual forb

Hemizonia congesta ssp. tracyi Asteraceae WI 1.8 5.25 3 2.0 0.7 0.4 4 1 Dicot Annual forbHemizonia halliana Asteraceae SI 3.0 12 4 3.0 2.4 1.2 1 Dicot Annual forbHesperevax sparsiflora var.

sparsifloraAsteraceae WI 1.8 7.25 4 1.5 1.6 0.8 1 1 1 Dicot Annual forb

Heterotheca oregona var.oregona

Asteraceae WI 2.0 6 3 3.0 1.7 1.0 1 1 1 Dicot Perennial forb

Hieracium bolanderi Asteraceae BE/SI 3.8 15 4 4.5 2.6 1.3 1 1 Dicot Perennial forbHieracium greenei Asteraceae WI 2.2 6.5 3 3.0 1.4 0.8 1 1 Dicot Perennial forbLagophylla glandulosa Asteraceae WI 2.0 6.1 3 3.0 1.7 1.0 1 1 Dicot Annual forbLagophylla minor Asteraceae BE 4.7 23.5 5 5.0 1.7 0.7 1 1 Dicot Annual forbLayia discoidea Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Annual forbLayia jonesii Asteraceae BE/SI 3.5 10.5 3 3.0 0.6 0.3 1b 1 Dicot Annual forbLayia septentrionalis Asteraceae SI 3.2 19 6 3.5 1.4 0.6 1b 1 Dicot Annual forbLessingia arachnoidea Asteraceae SE 6.2 18.5 3 6.0 0.0 0.0 1b 1 Dicot Annual forbLessingia filaginifolia var.

californicaAsteraceae WI/IN 1.3 4 3 1.0 1.5 0.9 1 1 1 1 1 Dicot Perennial

forb, ShrubLessingia hololeuca Asteraceae SI 2.5 7.5 3 3.0 1.2 0.7 3 1 1 Dicot Annual forbLessingia micradenia var.

glabrataAsteraceae BE 5.1 30.5 6 5.5 1.3 0.5 1b 1 Dicot Annual forb

Lessingia micradenia var.micradenia

Asteraceae BE 5.3 31.5 6 5.5 1.0 0.4 1b 1 Dicot Annual forb

Lessingia nemaclada Asteraceae WI 2.0 6 3 2.0 1.0 0.6 1 1 1 1 1 Dicot Annual forbLessingia occidentalis Asteraceae BE/SI 4.1 16.5 4 4.0 1.8 0.9 4 1 Dicot Annual forbLessingia ramulosa Asteraceae BE 5.4 27 5 6.0 1.3 0.6 1 1 Dicot Annual forbLuina hypoleuca Asteraceae WI/IN 1.4 4.25 3 2.0 1.0 0.6 1 1 1 Dicot Perennial forbMadia doris-nilesiae (5

Harmonia d.)Asteraceae BE 5.4 32.5 6 5.5 0.8 0.3 1b 1 Dicot Annual forb

Madia exigua Asteraceae WI 1.8 7.25 4 2.0 1.4 0.7 1 1 1 1 1 Dicot Annual forbMadia hallii (5 Harmonia h.) Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Annual forbMadia stebbinsii

(5 Harmonia s.)Asteraceae SE 6.1 42.5 7 6.0 0.0 0.0 1b 1 Dicot Annual forb

Malacothrix clevelandii Asteraceae SI 3.0 9.1 3 3.0 3.0 1.7 1 1 1 1 1 Dicot Annual forbMalacothrix floccifera Asteraceae WI 2.1 6.25 3 3.0 1.6 0.9 1 1 1 1 1 Dicot Annual forbMicropus amphibolus Asteraceae WI 2.4 7.25 3 1.0 3.1 1.8 1 1 1 Dicot Annual forbMicroseris douglasii Asteraceae WI/IN 1.3 4 3 1.0 0.6 0.3 1 1 1 1 Dicot Annual forbMonolopia gracilens Asteraceae WI 2.4 4.75 2 2.4 2.3 1.6 1 1 Dicot Annual forbPentachaeta bellidiflora Asteraceae WI 2.4 7.25 3 3.0 1.3 0.8 1b 1 Dicot Annual forb

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Pyrrocoma racemosa var.congesta

Asteraceae SE 6.2 18.5 3 6.0 0.0 0.0 2 1 Dicot Perennial forb

Pyrrocoma racemosa var.pinetorum

Asteraceae BE/SI 4.0 16 4 4.5 2.4 1.2 1 Dicot Perennial forb

Pyrrocoma racemosa var.racemosa

Asteraceae WI 1.7 5 3 1.0 2.1 1.2 1 1 1 Dicot Perennial forb

Raillardella pringlei Asteraceae SE 6.0 30 5 6.0 0.0 0.0 1b 1 Dicot Perennial forbRigiopappus leptocladus Asteraceae WI 1.9 7.5 4 2.0 1.3 0.7 1 1 1 1 1 Dicot Annual forbRudbeckia californica var.

glaucaAsteraceae BE 5.3 21 4 6.0 1.5 0.8 1 1 Dicot Perennial forb

Senecio clevelandii var.clevelandii (5 Packerac. v. c.)

Asteraceae SE 5.8 46.5 8 6.0 0.7 0.3 4 1 Dicot Perennial forb

Senecio clevelandii var.heterophyllus (5 Packera c.v. h.)

Asteraceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Perennial forb

Senecio eurycephalus var.eurycephalus (5 Packeraeurycephala var.eurycephala)

Asteraceae BE/SI 3.8 15 4 3.0 1.5 0.8 1 1 Dicot Perennial forb

Senecio eurycephalus var.lewisrosei (5 Packeraeurycephala var. lewisrosei)

Asteraceae SE 5.8 40.5 7 6.0 0.8 0.3 1b 1 Dicot Perennial forb

Senecio greenei (5 Packera g.) Asteraceae BE 5.3 32 6 6.0 1.6 0.7 1 1 Dicot Perennial forbSenecio layneae (5 Packera l.) Asteraceae BE 4.9 29.5 6 5.0 1.3 0.5 1b 1 Dicot Perennial forbSenecio macounii

(5 Packera m.)Asteraceae BE 5.1 20.5 4 6.0 2.0 1.0 4 1 Dicot Perennial forb

Solidago guiradonis Asteraceae SE 6.2 18.5 3 6.0 0.0 0.0 4 1 Dicot Perennial forbSolidago multiradiata Asteraceae WI/IN 1.1 2.1 2 1.1 1.3 1.0 1 1 Dicot Perennial forbStebbinsoseris decipiens Asteraceae WI 1.8 5.5 3 2.0 0.6 0.3 1b 1 1 Dicot Annual forbWyethia bolanderi Asteraceae WI 1.5 3 2 1.5 0.7 0.5 1 Dicot Perennial forbBerberis aquifolium var.

aquifoliumBerberidaceae WI 1.6 4.75 3 1.0 1.2 0.7 1 1 1 Dicot Shrub

Berberis aquifolium var. repens Berberidaceae WI 1.7 5 3 1.0 1.2 0.7 1 1 1 Dicot ShrubVancouveria chrysantha Berberidaceae SE 6.2 18.5 3 6.0 0.0 0.0 4 1 1 Dicot Perennial forb

(rhiz.)Vancouveria planipetala Berberidaceae WI 1.7 5 3 1.0 1.2 0.7 1 1 1 1 Dicot Perennial forb

(rhiz.)Cryptantha clevelandii var.

dissitaBoraginaceae BE/SI 4.4 17.5 4 4.5 2.1 1.0 1b 1 Dicot Annual forb

Cryptantha excavata Boraginaceae WI 1.5 3 2 1.5 2.1 1.5 1 Dicot Annual forb

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Cryptantha flaccida Boraginaceae WI 1.6 4.75 3 2.0 0.7 0.4 1 1 1 1 1 Dicot Annual forbCryptantha hispidula Boraginaceae SE 6.0 24 4 6.0 0.0 0.0 1 1 Dicot Annual forbCryptantha intermedia Boraginaceae WI/IN 1.4 4.1 3 1.0 1.5 0.9 1 1 1 1 1 Dicot Annual forbCryptantha mariposae Boraginaceae SE 6.2 18.5 3 6.0 0.0 0.0 1b 1 Dicot Annual forbCryptantha milobakeri Boraginaceae SI 3.3 9.75 3 3.0 2.6 1.5 1 1 1 Dicot Annual forbCryptantha sobolifera Boraginaceae WI/IN 1.4 4.1 3 1.0 1.5 0.9 1 1 Dicot Perennial forbHackelia bella Boraginaceae WI/IN 1.3 4 3 1.0 1.5 0.9 1 1 Dicot Perennial forbPectocarya pusilla Boraginaceae WI/IN 1.3 2.5 2 1.3 1.1 0.8 1 1 1 1 1 Dicot Annual forbArabis aculeolata Brassicaceae SE 6.1 24.5 4 6.0 0.0 0.0 2 1 Dicot Perennial forbArabis constancei Brassicaceae SE 5.9 41.5 7 6.0 0.4 0.1 1b 1 Dicot Perennial forbArabis koehleri var. stipitata Brassicaceae SE 5.5 16.5 3 6.0 1.2 0.7 1b 1 Dicot Perennial forbArabis macdonaldiana Brassicaceae BE 5.4 32.5 6 6.0 1.0 0.4 1b 1 1 Dicot Perennial forbArabis oregana Brassicaceae BE/SI 3.8 11.5 3 3.0 2.1 1.2 4 1 1 Dicot Perennial forbArabis subpinnatifida Brassicaceae SI 3.2 16 5 3.0 1.6 0.7 1 1 Dicot Perennial forbArabis suffrutescens var.

horizontalisBrassicaceae BE/SI 3.9 27 7 3.0 1.6 0.6 1 1 Dicot Perennial forb

Arabis suffrutescens var.suffrutescens

Brassicaceae SI 2.9 17.5 6 2.5 1.9 0.8 1 1 Dicot Perennial forb

Cardamine californica var.cuneata

Brassicaceae WI 1.9 3.75 2 1.9 1.6 1.1 1 Dicot Perennial forb(rhiz.)

Cardamine nuttallii var.gemmata

Brassicaceae BE 5.2 15.5 3 5.0 1.0 0.6 1b 1 Dicot Perennial forb(rhiz.)

Cardamine pachystigma var.dissectifolia

Brassicaceae BE 5.4 48.5 9 6.0 1.0 0.3 3 1 1 Dicot Perennial forb(rhiz.)

Cardamine pachystigma var.pachystigma

Brassicaceae WI 2.0 6 3 2.0 1.0 0.6 1 1 Dicot Perennial forb(rhiz.)

Caulanthus amplexicaulis var.barbarae

Brassicaceae SE 6.2 18.5 3 6.0 0.0 0.0 1b 1 Dicot Annual forb

Draba aureola Brassicaceae SI 2.7 8 3 3.0 0.6 0.3 1b 1 Dicot Perennial forbDraba carnosula Brassicaceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Perennial forbDraba howellii Brassicaceae WI/IN 1.4 4.25 3 1.0 1.4 0.8 1 Dicot Perennial forbErysimum franciscanum Brassicaceae SI 3.0 9 3 3.0 0.0 0.0 4 1 1 1 Dicot Perennial forbGuillenia flavescens Brassicaceae WI 2.3 9.25 4 2.5 1.6 0.8 1 1 1 Dicot Annual forbStreptanthus albidus ssp.

albidusBrassicaceae BE 5.3 31.5 6 6.0 1.3 0.5 1b 1 1 Dicot Annual forb

Streptanthus albidus ssp.peramoenus

Brassicaceae BE/SI 4.3 34.5 8 4.5 1.9 0.7 1b 1 1 Dicot Annual forb

Streptanthus barbatus Brassicaceae SE 5.6 28 5 6.0 0.5 0.2 1 Dicot Perennial forbStreptanthus barbiger Brassicaceae SE 6.0 24 4 6.0 0.0 0.0 4 1 Dicot Annual forbStreptanthus batrachopus Brassicaceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 1 Dicot Annual forb

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Streptanthus brachiatus var.brachiatus

Brassicaceae SE 5.6 22.5 4 6.0 1.0 0.5 1b 1 Dicot Annual,Perennialforb

Streptanthus brachiatus var.hoffmanii


Streptanthus breweri var.breweri

Brassicaceae SE 5.7 40 7 6.0 0.8 0.3 1 1 1 1 Dicot Annual forb

Streptanthus breweri var.hesperidus

Brassicaceae SE 6.1 24.5 4 6.0 0.0 0.0 1b 1 Dicot Annual forb

Streptanthus drepanoides Brassicaceae SE 6.1 36.5 6 6.0 0.0 0.0 4 1 1 1 Dicot Annual forbStreptanthus glandulosus ssp.

glandulosusBrassicaceae WI 1.9 5.75 3 2.0 1.1 0.7 1 1 1 Dicot Annual forb

Streptanthus glandulosus ssp.pulchellus

Brassicaceae BE 4.9 24.5 5 6.0 1.8 0.8 1b 1 1 Dicot Annual forb

Streptanthus glandulosus ssp.secundus

Brassicaceae SI 3.3 20 6 3.0 1.5 0.6 1 1 Dicot Annual forb

Streptanthus glandulosus ssp.secundus var. hoffmanii

Brassicaceae SI 3.0 3 1 3.0 — — 1b 1 Dicot Annual forb

Streptanthus howellii Brassicaceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 Dicot Perennial forbStreptanthus insignis ssp.

insignisBrassicaceae BE/SI 4.0 20 5 4.0 2.4 1.1 1 Dicot Annual forb

Streptanthus insignis ssp. lyonii Brassicaceae SI 3.3 16.5 5 2.0 2.7 1.2 1b 1 Dicot Annual forbStreptanthus morrisonii ssp.

elatusBrassicaceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 Dicot Annual,

Perennialforb

Streptanthus morrisonii ssp.hirtiflorus


Streptanthus morrisonii ssp.kruckebergii


Streptanthus morrisonii ssp.morrisonii


Streptanthus niger Brassicaceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 1 Dicot Annual forbStreptanthus polygaloides Brassicaceae SE 5.7 28.5 5 6.0 0.9 0.4 1 Dicot Annual forbStreptanthus tortuosus var.

suffrutescensBrassicaceae WI 1.6 8.2 5 2.0 1.6 0.7 1 1 1 Dicot Annual,

Perennialforb

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Streptanthus tortuosus var.tortuosus

Brassicaceae WI/IN 1.4 4.25 3 2.0 1.0 0.6 1 1 1 Dicot Annual,Perennialforb

Thelypodium brachycarpum Brassicaceae SI 3.3 10 3 3.0 0.6 0.3 4 1 1 Dicot Annual,Perennialforb

Thlaspi californicum Brassicaceae SE 6.1 30.5 5 6.0 0.0 0.0 1b 1 Dicot Perennial forbThlaspi montanum var.

montanumBrassicaceae BE/SI 4.4 22 5 4.0 1.5 0.7 1 1 Dicot Perennial forb

Campanula angustiflora Campanulaceae BE/SI 3.9 19.25 5 4.0 2.4 1.1 1 1 Dicot Annual forbCampanula exigua Campanulaceae BE/SI 3.9 19.5 5 4.0 1.5 0.7 1b 1 1 Dicot Annual forbCampanula griffinii Campanulaceae SE 6.0 18 3 6.0 0.0 0.0 1 1 1 Dicot Annual forbCampanula rotundifolia Campanulaceae BE 5.0 15 3 6.0 1.7 1.0 1 Dicot Perennial forbCampanula scabrella Campanulaceae SI 2.5 10 4 2.5 1.3 0.6 4 1 Dicot Perennial forb

(rhiz.)Campanu

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