E A GE E A OG AM
DEMOGRAPHY OF CAU A US CA I O ICUS, EM E IA CO G O II, AND E IAS UM OO E , AND VEGETATION CHARACTERISTICS OF ENDANGERED SPECIES POPULATIONS IN THE SOUTHERN SAN JOAQUIN VALLEY AND THE CARRIZO PLAIN NATURAL AREA IN 1993
FINAL REPORT
SUBMITTED BY:
Ellen Cypher San Joaquin Valley Endangered Species ey Recovery Planning Program 2727 North Grove Industrial Drive, Suite 125 Co ( o51 Fresno, CA 93727 (209) 453-1103 1 1
SUBMITTED TO:
Deborah Hillyard California Department of Fish and Game Natural Heritage Division 1416 9th Street Sacramento, CA 95814-2090
FUNDED BY:
Emergency Drought Relief Project Contract No. CA HER 031293
23 May 1994 • ABSTRACT
Populations of two endangered and one threatened plant species were studied on the Carrizo Plain Natural Area (CPNA) and in the San Joaquin Valley of California in order to provide baseline data for continuing research on their ecology and responses to drought and site management. Results will be incorporated into a multi-species recovery plan as well as restoration and management plans for the CPNA and other preserves.
In terms of plant size and reproduction, the most successful populations of Caulanthus caqomicus were those on the CPNA, whereas Lembertia congdonii was equally successful on the CPNA and in the Kettleman Hills. Of the four Eriastrum hooveri sites studied, plants on the CPNA were smallest with poorest reproduction.
Grazing effects were compared in three populations of L. congdonii, each of which was divided into grazed and ungrazed portions. Reproduction of L. congdonii was greater in grazed than in ungrazed areas of the Carrizo Plain and Kettleman Hills, whereas the reverse was found on the Elkhorn Plain.
Distribution and morphological characters of C. californicus and L. congdonii were evaluated relative to giant kangaroo rat (Dipodomys ingens) precincts on the Carrizo and Elkhorn Plains. Although C. ca4omicus occurred more frequently on precincts than did random points, plants growing on precincts did not differ significantly in size or reproduction from those growing between precincts. Conversely, L. congdonii occurred randomly with respect to precincts. The two L. congdonii populations studied differed in plant response to giant kangaroo rat precincts in terms of plant size and flower head production, but not in survival and achene production.
Habitat characteristics of the listed plant and animal species were assessed on 19 vegetation transects on the CPNA and 43 in the San Joaquin Valley. Exotic plants dominated in cover at most sites, yet exotics accounted for less than one-third of species composition, on average. Plant diversity was high on the CPNA compared to other areas sampled, and species richness was greater in ungrazed than in grazed areas. Native plants, including the listed species, apparently persisted in the seed bank during the recent drought.
Reintroduction of Caulanthus cal&micus, Lembertia congdonii, and Eriastrum hooveri is not necessary on the CPNA at this time. Exotic plants are not likely to be eradicated from the area, but active management (e.g., planting, seeding, burning) could increase the proportion and distribution of native plants, particularly shrubs and perennial grasses. Controlled grazing studies are a prerequisite to management decisions. Additional research is necessary to determine the extent of the existing seed bank and the appropriate management techniques for enhancement of rare plant populations and rehabilitation of plant communities affected by drought and historic agricultural practices. ACKNOWLEDGMENTS
We appreciate the efforts of the many individuals who provided field assistance, advice, and logistical support during the 1993 field season. Wendy Stevens and Kirk Lenington were primarily responsible for data collection on the Carrizo Plain Natural Area. Field work in other study areas was conducted by Graciela Melgoza Hinshaw, Carol Ashbrook, Norman Vulich, and Shirley May. Terrence Trasatti, Heather Johnson, Michael Allenger, Mark McFall, and Brian Cypher provided assistance at various sites. Rebecca Roth provided invaluable logistical support. Many employees of the U.S. Bureau of Land Management directed us to suitable study sites, obtained permission for us to cross private land, helped collect data, and provided historical information; Russ Lewis and Bruce Delgado were particularly helpful in this regard. We also thank Tony Nelson of The Nature Conservancy for allowing us to use facilities on the Carrizo Plain.
Funding for this research was provided by the following agencies: (1) California Department of Fish and Game, Natural Heritage Division, Emergency Drought Relief Project, (2) U.S. Bureau of Land Management., Bakersfield District, (3) U.S. Bureau of Reclamation, Friant Division, and (4) U.S. Fish and Wildlife Service, Sacramento Field Office.
ii TABLE OF CONTENTS
ABSTRACT
ACKNOWLEDGMENTS ii
LIST OF TABLES
LIST OF FIGURES vi
INTRODUCTION AND PROJECT OBJECTIVES 1
METHODS 1 Demography 1 Population Trends 3 Grazing Effects 3 Endangered Plant-Giant Kangaroo Rat Associations 4 Habitat Characterization 4 Data Analyses 5 Coordination with Dr. Paula Schiffman 5
RESULTS Cau a us ca o cus E ias um oo e i 9 em e ia co g o ii 15 Habitat Characterization 21
DISCUSSION 26
RECOMMENDATIONS 32
LITERATURE CITED 33
KNOWLEDGEABLE INDIVIDUALS 37
APPENDICES
A. Names, numbers, and locations of study sites in the San Joaquin Valley, California, and adjacent areas that were used for demographic monitoring of listed plant species in Spring 1993 39
B. Locations of vegetation transects sampled in Spring 1993 in the San Joaquin Valley and adjacent foothills, California 41
iii C. Cover, species composition, and diversity on vegetation transects sampled in the San Joaquin Valley and adjacent foothills, California, Spring 1993 45
D. Photographs of vegetation transects from the Carrizo Plain Natural Area, San Luis Obispo County, California 50
iv LIST OF TABLES
1. Comparison of attributes among four Caulanthus ca4onficus populations .... 8
2. Comparison of Caulanthus caqomicus attributes on vs. off giant kangaroo rat precincts at Site 1 on the Carrizo Plain, San Luis Obispo County, California 10
3. Comparison of Caulanthus californicus attributes on vs. off giant kangaroo rat precincts at Site 2 on the Carrizo Plain, San Luis Obispo County, California 11
4. Results of nonparametric analysis of variance for Caulanthus cagomicus at Carrizo Plain Sites 1 and 2, on and off giant kangaroo rat precincts 12
5. Comparison of attributes among four Eriastrum hooveri populations 13
6. Comparison of Eriastrum hooveri attributes in grazed vs. ungrazed areas at Site 8 in the Kettleman Hills, Kings County, California 14
7. Comparison of attributes among seven Lembertia congdonii populations in Spring 1993 16
8. Comparison of Lembertia congdonii attributes in grazed vs. ungrazed areas at Site 4 on the Carrizo Plain, San Luis Obispo County, California 18
9. Comparison of Lembertia congdonii attributes in grazed vs. ungrazed areas at Site 6 on the Elkhorn Plain San Luis Obispo County, California 19
10. Comparison of Lembertia congdonii attributes in grazed vs. ungrazed areas at Site 8 in the Kettleman Hills, Kings County, California 20
11. Comparison of Lembertia congdonii attributes on vs. off giant kangaroo rat precincts at Site 4 on the Carrizo Plain, San Luis Obispo County, California 22
12. Comparison of Lembertia congdonii attributes on vs. off giant kangaroo rat precincts at Site 6 on the Elkhorn Plain, San Luis Obispo County, California 23
13. Results of nonparametric analysis of variance for Lembertia congdonii at Sites 4 and 6 on the Carrizo and Elkhorn Plains, on and off giant kangaroo rat precincts, in grazed and ungrazed areas 24 14. Cover and species composition of native versus exotic plant species on vegetation transects in habitats of listed plant or animal species in the southern San Joaquin Valley and adjacent areas, Spring 1993 25
15. Mean cover and species composition of native versus exotic plant species on vegetation transects from various plant communities and treatments sampled on the Carrizo Plain Natural Area, San Luis Obispo County, California, in Spring 1993 27
16. Plant species most commonly encountered on 19 transects in endangered species populations on the Carrizo Plain Natural Area, San Luis Obispo County, California, in Spring 1993 28
vi LIST OF FIGURES
1. Approximate locations of study sites in the San Joaquin Valley, California, and adjacent areas that were used for demographic monitoring of listed plant species in Spring 1993 2
2. Approximate locations of vegetation transects sampled in Spring 1993 in the San Joaquin Valley and adjacent areas, California 6
vii INTRODUCTION AND PROJECT OBJECTIVES
Although Cau a us ca o icus (S. Watson) Payson and em e ia co g o ii (Gray) E. Greene are federally listed as endangered species, and E ias um oo e i (Jepson) H. Mason is federally listed as threatened (U.S. Department of the Interior, Fish and Wildlife Service 1990), little is known about the biology, demography, and management needs of these plant species. Field studies in the San Joaquin Valley of California and adjacent areas were conducted in Spring 1993 in order to (1) monitor the survival, growth, and reproduction of these species in various parts of their ranges and under different grazing conditions, (2) determine baseline population size so population trends can be correlated with future weather cycles and grazing regimes, (3) evaluate the association of endangered plants with the endangered giant kangaroo rat ( i o omys i ge s , and (4) describe the habitat of listed plant and animal species. Information on the relative performance of each species under the observed grazing regimes, the role of giant kangaroo rats in rare plant growth and reproduction, and the effect of plant community composition on species occurrence will be incorporated into a comprehensive recovery plan. Additionally, data from the Carrizo Plain Natural Area (CPNA) in San Luis Obispo County, California, will be valuable for determining strategies for restoring native vegetation in that area, which has been degraded due to farming and ranching operations over many decades, and more recently, due to drought.
METHODS
Demography
Demographic studies were conducted at 6 sites on the CPNA and 8 sites throughout the San Joaquin Valley (Figure 1, Appendix A). Populations of the listed plants were chosen for monitoring from among known localities based on (1) avoiding conflict with other research programs, (2) opportunity for comparison of different site conditions (e.g., cattle grazing, occurrence of giant kangaroo rats), (3) distribution throughout the known range of each species, and (4) site accessibility.
Individual plants for study were selected randomly in one of two ways, depending on overall population size. In populations totalling 250 individuals or fewer (Cau a us caqomicus at Sites 10 and 11, em e ia co g o ii at Sites 9 and 14), all plants were numbered individually, then 50 were chosen with the aid of a random number table. At the remaining sites, plants were located by pacing random distances along transects spaced 10 m apart within the population, then the closest individual of the target species to the random point was located. Circular 0.25 m 2 sampling plots were established that included the closest individual and any others within a 0.28 m radius. In plots containing more than three individuals, three were selected randomly and marked for intensive monitoring.
Few individuals of C. ca O icus were found off-precinct at Site 1 by the random process described above. Therefore, a modified method was used to select plants for Ca a • ‘ Q o • a[ -"C A cu -•- W ge u e$1 A
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Data collected biweekly on each marked individual included rosette diameter (in Cau a us ca o icus , maximum stem height/length; number of stems, buds, flowers, and fruits; fruit length (C. c4o icus and number of seeds/carpels per mature fruit (C. ca o icus and E. oo e i or number of mature achenes per head ( . co g o ii . Causes of mortality and/or damage to plant parts also were noted at each visit. Damage was attributed to cattle, insects, and/or kangaroo rats if signs indicated their presence (e.g., cattle hoof prints or droppings, small holes chewed by insects, kangaroo rat scats or burrows); otherwise, damage was recorded as due to unknown causes. Although all of the intensively-studied species were annuals and were therefore expected to die by the end of the growing season, marked individuals were considered to have survived if they persisted long enough to set seed.
Population Trends
Long-term monitoring will allow population sizes to be correlated with annual environmental variation and habitat management. Total counts were obtained in populations of C. ca O icus used for demographic studies. However, populations of E. oo e i and . co g o ii were too large for complete censuses, so density was estimated instead. Density was estimated by counting individuals of the target species in a 50 x 1 m belt centered on the vegetation transect (see "Habitat Characterization"). In extremely dense populations, one subsample was obtained per square meter in randomly- placed plots measuring 0.5 m x 0.2 m. Statistical analyses were not conducted on densities.
Grazing Effects
Time constraints prohibited controlled grazing studies in Spring 1993. Instead, demographic monitoring was carried out in several populations that were divided into grazed and ungrazed portions by fences; plant size and demographic variables then were compared within populations with respect to grazing. Two sites on the CPNA (#4 and #6) that were subject to cattle grazing from February through April 1993 met this criterion, and both supported . co g o ii. A third such site in the Kettleman Hills (#8) included both E. oo e i and . co g o ii. Known populations of C. ca o icus were not grazed during the flowering season in 1993. 4 Endangered Plant-Giant Kangaroo Rat Associations
Associations between Cau a us ca o icus or em e ia co g o ii and giant kangaroo rats were investigated at Sites 1, 2, and 4 on the Carrizo Plain, where the animals occur in dense colonies. Giant kangaroo rats modify their habitat extensively by burrowing in the soil, clipping vegetation, and collecting seed (Williams and Kilburn 1991). Their burrows are concentrated into roughly circular areas known as precincts (Grinnell 1932), whereas the areas between precincts are known as interspaces or "off- precinct." For the purposes of this study, the outer margin of a precinct was delimited by the openings of converging horizontal burrows and their associated ramps. The delimited area was conclusively determined to be a precinct if the surface exhibited mounding, openings of vertical burrows, and/or accumulations of seed. Interspaces were defined as undisturbed areas between burrows that led in opposite directions. Areas in the vicinity of isolated burrows that exhibited no characteristic signs of giant kangaroo rat activity were classified as unknown.
Frequency of occurrence of rare plant individuals on precincts was compared to that of random points to determine whether or not the plants occurred disproportionately on precincts. All individuals of C. ca , in the two monitored populations were inspected to determine if they were growing on precincts. Occurrence of . co g o ii on precincts was determined during the process of evaluating random points. Placement of random points was determined by selecting a number of paces from a random-number table, then pacing along transects that were spaced 10 m apart within the limits of the population. The number of random points was not determined in advance, but instead depended on the size of the random numbers relative to the extent of each population. Each random point was classified as occurring on a precinct or in an interspace. At Site 4, after a random point was identified, the nearest individual of . co g o ii was located and evaluated in terms of occurrence on a precinct.
Individual plants of both species were monitored on giant kangaroo rat precincts and in interspaces in conjunction with demographic studies to compare survival, size, and reproduction with respect to giant kangaroo rat activity.
Habitat Characterization
Plant communities supporting listed plant and animal species were characterized using the vegetation sampling procedure established by the California Native Plant Society (1993). Consistent with this method, point-intercepts spaced at 50-cm intervals along a 50-m transect were used to determine percent cover for each vascular plant species in each stratum. In addition, all plant species occurring within a 2.5-m belt along each side of the transect were recorded; site information such as elevation, slope and aspect were noted; and photographs were taken from each end of the transect. More than one transect was established in certain populations that varied in grazing treatment 5 or topography. At the CPNA, vegetation was sampled in populations of the three listed plants already mentioned, as well as in populations of i o omys i ge s and Gam e ia sila. Transects also were placed in populations of other federally-listed plant and animal species in the San Joaquin Valley and adjacent foothills (Figure 2, Appendix B).
Plant community type was determined for each transect based on dominant species and site location (Holland 1986). Species identifications were determined using the Jepson Manual (Hickman 1993) and local floras (Twisselmann 1956, Hoover 1970).
Data Aaa yso
Nonparametric statistics were used to compare quantitative data because the variables were not normally distributed and neither variances nor sample sizes were equal. Single-factor comparisons employed the Mann-Whitney test for paired samples and the Kruskal-Wallis test for three or more samples; the resulting statistics were then converted to an approximate Chi-square value. For simultaneous analyses of two or more factors, raw data were ranked and then standard analysis of variance techniques were performed on the ranks (Zar 1984). Linear regression was not attempted because the data did not meet the necessary assumptions, even after transformation.
Contingency table analysis employing a Chi-square statistic was used to compare frequency data such as survival or occurrence on precincts; 2 x 2 contingency table analysis included Yates correction for continuity. Correlations between variables were calculated using Spearman 's rank correlation coefficient, a nonparametric technique. Maximum values of diameter, height, and number of stems for each individual were used in statistical analyses. Total flower production was estimated by summing the number of buds, flowers, and fruits on each plant at each visit; the maximum value per plant was used for statistical analyses.
The Shannon Diversity Index was calculated for each vegetation transect using natural logarithms (Brower and Zar 1984, Zar 1984). Because this index incorporates both species richness and abundance, only those species that were intercepted on the transect were included in the calculations. Data were pooled over all strata for each transect. Cover values were relativized for analyses by transect, but raw (absolute) cover values were used for comparisons by plant species.
Coordination with Dr. Paula Schiffman
In order to ensure consistency, standardized vegetation sampling techniques were agreed upon and practiced jointly prior to data collection. Moreover, to maximize data collection and avoid overlap, the two research groups focused on different locations within the CPNA. Employees of the San Joaquin Valley Endangered Species Recovery Figure 2. Approximate locations of vegetation transects sampled in Spring 1993 in the San Joaquin Valley and adjacent areas, California. 7 Planning Program concentrated on sampling in known populations of listed plant and animal species, whereas Dr. Schiffman directed her sampling to common vegetation associations. However endangered animals, particularly i o omys i ge s, may have been present in the latter areas. Finally, copies of all transect data sheets and maps, as well as a computer file on diskette, were provided to Dr. Schiffman after the field season had ended.
ESU S
Data collection began on 15 February 1993, when the C. caqomicus and . co g o ii populations on the Carrizo and Elkhorn Plains were in the seedling and/or rosette stages, and E. oo e i seeds had not yet germinated. Monitoring continued through senescence of all marked individuals. The last day of data collection was for monitoring of E. oo e i on 8 June 1993. Data on rare plant occurrence on giant kangaroo rat precincts were collected from 8 April through 10 May 1993.
Factors responsible for mortality were not readily apparent in most monitored populations. Typically, plants were alive at one visit and dead or missing on subsequent visits. Primary causes of damage to marked plants were noted when known, but these agents may or may not have contributed to plant mortality. A combination of age, heat, and water stress most likely was responsible for the death of many plants.
Cau a us ca i omicus
Plants from the two Carrizo Plains populations were significantly greater in most measurements of size and fecundity than those from the Kreyenhagen Hills (Table 1). When all four populations were compared, the two from the Carrizo Plain differed significantly from each other in rosette diameter, number of stems, and total flower production. The two Kreyenhagen Hills populations differed from each other in plant height and flower production. Competition from other herbs may not have been related to successful growth and reproduction of C. ca omkus because the herbaceous cover was similar on three sites that encompassed the range of plant size and fruit production.
Plants on the CPNA were subject to burial under loose soil and/or stem clipping by giant kangaroo rats, but damaged C. ca i o icus individuals typically produced new shoots from the rosette. Plants in the Kreyenhagen Hills also showed damage to the stems and leaves, but the source could not be determined. Flowering and fruiting in the Kreyenhagen Hills occurred several weeks earlier than on the Carrizo Plain, which is approximately 300 m higher in elevation.
Table 1. Comparison of attributes among four Cau a us ca i o icus populations (see Appendix A for exact locations). Means are shown ± standard error. Sample size is as indicated at the top of each column unless given in parentheses.
O U A IO CA I O AI K EYE AGE I S 1 1 11 ( = ( = 5 ( = 5 ( = 5 um e su i i g o ui i g / 37/5 /A /A ( 3 ( 1 gia 5/ 7/5 /5 ° /5 ° ka ga oo a s (37 9 ( & ( ( um e i sec s 3/ /5 /5 /5 amage y ( 5 (1 7 ( (3 u k ow / /5 7/5 ° 5/5 ° sou ces ( ( (1 (9 ose e iame e (cm 15 555° 1 35 ± 7 ± 9 ± 35 ± 1 eig (cm 99 1 5k° 1 9 15 1 ± 1 9 ± 1 1 ± ± 75 um e o s ems 3 19 ° 1 e 1 15 ± 35 ± 5 ± 15 ± 1 o a owe o uc io 73 3 3 5 1 1 39° ± 19 1 ± 9 7 ± 1 7 ±135 ui e g (cm 3 71 3 1 1 1° 5° ± 1 ± 1 ± 3 ± 1 ( = 1 9 ( = 3 ( = 0 ( = 3 um e o see s e ui 33 753 &go 1 ± 3 ± 3 ± 1 7 ( = 1 9 ( = 3 ( = 0 ( = 3 a e owe s i s o se e i 1 Ma c 1 Ma c (9 Ma c (9 Ma c 1993
a e ma u e ui s i s 7 A i 7 A i (9 Ma c (9 Ma c o se e i 1993 o u a io si e 9 1 9 5 197 e aceous co e ( 9 5 9 95-97 a Means followed by the same letter are not significantly different at = 0.05. Not calculated due to missing data. 2 Flowering and/or fruiting well advanced at date of first visit. 9 Caulanthus ca4ornicus individuals were significantly associated with giant kangaroo rat precincts on the Carrizo Plain (Tables 2 and 3). A number of plants at Site 2 were omitted from the analysis because their occurrence on or off precincts could not be determined with certainty. Giant kangaroo rats were not present at the Kreyenhagen Hills sites. Few attributes differed significantly on versus off precincts in either Carrizo Plain population (Tables 2 and 3). Rosette diameter at Site 2 was significantly greater off precinct, and at a = 0.10, survival of C. cal&micus individuals at Site 1 was significantly higher on precincts than off. However, clipping of plant parts by giant kangaroo rats did not differ significantly with respect to precincts. Variability between sites accounted for the majority of differences between the two populations (Table 4). When differences due to occurrence on precincts were factored out, fruit length and number of seeds per fruit differed significantly between Sites 1 and 2 (Table 4), which had not been evident when all four populations were compared without respect to precincts (Table 1). Site-by-precinct interactions were not significant, with the possible exception of rosette diameter (Table 4).
Eriastrum hooveri
Survival, height, and flower production in E. hooveri were significantly lower on the Carrizo Plain than at all of the other sites (Table 5). Virtually all of the plants at all sites had a single stem, although some individuals did branch. Therefore, data for number of stems are not presented. Eriastrum hooveri plants did not resprout following stem breakage due to trampling or herbivory by cattle, rodents, or insects. However, physical damage and competition from species other than Eriastrum probably were not responsible for the high mortality rate at the Carrizo Plain site because both were comparatively low. Instead, mortality at Site 3 may have been primarily density- dependent. Fresno, Kern, and Kings County populations flowered and produced fruit well in advance of the population in San Luis Obispo County. The elevation at Site 3 is 400-600 m higher than at the other E. hooveri study sites.
Several problems were encountered at Site 3, which contained a mixture of three Eriastrum species. The final sample size was small because fewer than half of the marked individuals proved to be E. hooveri. In addition, the density estimate was invalid because it was obtained prior to flowering and may have included stems of other species. Similarly, Site 13 included both E. hooveri and E. pluriflorum. However, data collection was not compromised because no individuals of the latter species had been marked, and density was estimated while the plants were in flower.
At Kettleman Hills Site 8, where E. hooveri was monitored in both grazed and ungrazed areas, no significant differences in survival, size, or reproduction were found at the 0.05 level (Table 6). However, the data suggest that (1) survival was higher in the grazed area, (2) plants were taller in the ungrazed area, and (3) fecundity did not differ due to grazing. Sources of damage differed significantly between the grazed and 1
a e Com a iso o Caulazuhus califomicus a i u es o s o gia ka ga oo a eci c s a Si e 1 o e Ca i o ai Sa uis O is o Cou y Ca i o ia (see A e i A o e ac oca io Mea s a e s ow ± s a a e o Sam e si e is as i ica e a e o o eac co um u ess gi e i a e eses
OCA IO E A I E O O 2 O ECI C ( = ( = 15 um e su i i g o ui i g 31/ /15 3 3 7 (7 5 ( gia 15/ 7/15 31 575 um e kangaroo rats (3 1 ( 7 amage y i sec s 1/ 1/15 9 9 ( 3 ( 7 ose e iame e (cm 7 ± 9 35 ± 1 7 7 7 7 eig (cm 5 ± 3 31 ± 3 9 13 um e o s ems 3 ± 7 3 53 ± 17 7 o a owe o uc io 1 57 ± 7 51 7 ± 9 37 ui e g (cm 3 57 ± 1 3 7 ± 19 91 339 ( = (n = 3 um e o see s e ui 7 ± 3 ± 3 31 1 (n = (n = 3 C. ca i o icus / 9 / 9 Number of (5 3 ( 1 7 occurrences 5 1 a om 1 / / oi s (3 1 ( 1 9 1 1
a e 3 Com a iso o C,cui a us ca i o icus a i u es o s o gia ka ga oo a eci c s a Si e o e Ca i o ai Sa uis O is o Cou y Ca i o ia (see A e i A o e ac oca io Mea s a e s ow ± s a a e o Sam e si e is as i ica e a e o o eac co um u ess gi e i a e eses
OCA IO E A I E O O 2 O ECI C ( = ( = 7 um e su i i g o ui i g 1 / 19/ 7 15 9 ( 1 5 (7 gia 1 / 11/ 7 1 5 15 um e ka ga oo a s ( 1 5 ( 7 amage y i sec s 3/ 3/ 7 15 7 1 (113 (7 ose e iame e (cm 5 ± 55 7 ± 1 79 9 eig (cm 17 ± 3 19 73 ± 3 um e o s ems 1 1 ± 3 ± 5 o a owe o uc io 7 ± 1 5 ± 17 5 3 ui e g (cm 3 ± 19 3 ± 17 5 ( = 14) ( = 1 um e o see s e ui 7 ± 3 1 3 ± 3 3 7 (n = 1 ( = 1 um e o C. ca i o icus 7 /13 57 /13 occurrences (57 ( 9 a om 7/ 7 / 7 oi s ( 5 9 (7 1 1
Table 4. Results of nonparametric analysis of variance for Cau a u us ca i omicus at Carrizo Plain Sites 1 and 2, on and off giant kangaroo rat precincts. See Appendix A for exact site locations.
SI E SI E ECI C ECI C AC O I E AC IO ose e iame e 1 1 3 1 75 1 9 3 5 eig 1 1 1 7 3 5 1 5 177 um e o s ems 1 1 1 1 77 3 3 1 9 7 o a owe o uc io 1 1 5 3 7 1 9 1 57 1 ui e g 1 1 73 11 1 1 1 751 um e o see s e ui 1 1 1 15 3 5 1 9 151 13
Table 5. Comparison of attributes among four E ias um oo e i populations (see Appendix A for exact locations). Means are shown ± standard error. Sample size is as indicated at the top of each column unless given in parentheses.
O U A IO CA I O ACA I OS KE EMA OKE AI I S I S 13 3 7 ( = 13 ( = 7 ( = 55 ( = 5 um e su i i g o ui i g 7/ 7 1/55 1/ 5 75/13 ( 5 9 (7 5 ( 3 1 (57 7 ca e / 7 /55 1 / 5 /13 am i g ( (1 9 ( 7 7 (
um e o e s / 7 /55 13/ 5° 9/13 amage y ( ( ( ( 3 u k ow / 7 5/55° / 5 /13 sou ces (7 (9 1 (9 (3 1 eig (cm 5 3 5 15 5 1 ± 3 ± 37 ± 3 ± 3 o a owe o uc io 1 11 1 ± ± 1 ± 9 ± 1 37 um e o see s e ui 5 3 7 5a /A ± ± ± 9 ( = ( = ( = 1 a e owe s i s o se e i 3 A i 31 Ma c Ma c 5 A i 1993 a e ui s i s o se e i May 15 A i 13 A i 19 A i 1993
o u a io si e ca 1 5 1 ca 1 1
E. oo e i e si y ( a s/m ( 9 3 9 1 7 e aceous co e ( - 9 9 9-9 -9
I Means followed by the same letter are not significantly different at = 0.05. 1 Not calculated due to missing data Estimate from Lewis (1992). 3 Density estimate may include stems of E. a o um and E. i iesum. Three belts sampled within population. 1
a e Com a iso o Eriastrum hooveri a i u es i g a e s u g a e a eas a Si e i e Ke ema i s Ki gs Cou y Ca i o ia (see A e i A o e ac oca io Mea s a e s ow ± s a a e o Sam e si e is as i ica e a e o o eac co um u ess gi e i a e eses
SI E G A I G EGIME G A E U G A E I 1993 ( = 3 ( = 33 um e su i i g o ui i g /3 17/33 91 (75 (51 5 ca e 1 /3 /33 9 am i g (5 3 ( um e amage y o e s /3 13/33 13 39 ( (39 u k ow /3 /33 35 sou ces ( (1 eig (cm 53 ± 31 5 7 ± 9 3 5 5 o a owe o uc io 3 ± 137 3 ± 13 77 um e o see s e ui ± 33 3 ± 3 3 3 ( = 3 ( = E. oo e i e si y i e 1 7 1 • • a sec ( a s/m e aceous co e ( 9 9 -9 • •
wo e s sam e wi i u g a e o u a io • o es e 15 ungrazed areas, with the primary factors being cattle trampling and rodent clipping, respectively. Damage rates were higher than mortality rates because plants that were damaged after setting seed were considered to have survived to reproduction. The reduced vegetation cover in the grazed area may have contributed to the greater survival rates and shorter plants. Conversely, competition from the tall, dense herbaceous cover in the ungrazed area may have led to greater mortality of E. hooveri but taller growth of the surviving individuals. Density estimates were not instructive due to their extreme variability, even with respect to a given grazing treatment.
Lembertia congdonii
Lembertia congdonii stems grew both upright and prostrate; therefore, maximum stem length rather than height was used as an index of size. Significant damage to marked L. congdonii plants was caused by kangaroo rats (Dipodomys spp.), cattle, and unknown herbivores (Table 7). However, the plants typically produced additional stems to replace those lost to herbivory and/or trampling, and thus damage was not directly related to mortality. Of the five general areas where L. congdonii was studied, the Carrizo Plain, Elkhorn Plain, and Kettleman Hills all exhibited exceptional plant size and flower head production compared to the Jacalitos and Panoche Hills (Table 7). However, achene production at Site 8 in the Kettleman Hills significantly exceeded all other sites. Flowering and fruiting of plants in the northern populations preceded those in the southern populations because the former are several hundred meters lower in elevation. The number of individuals varied greatly among populations, with the fewest at Sites 7, 9, and 14. On the Carrizo and Elkhorn Plains, Lembertia congdonii was essentially continuous, with the combined populations representing millions of plants (Lewis 1993). Most L. congdonii populations senesced before density estimates could be obtained. Herbaceous cover was fairly consistent over all of the study sites. Heavy grazing may have reduced herbaceous cover as well as L. congdonii size and reproduction at Site 14 in the Panoche Hills.
Grazing did not affect performance of L. congdonii predictably in the divided populations studied (Tables 8-10). Survival did not differ significantly between grazed and ungrazed areas at Sites 4 and 6. Plants at Site 8 were marked too late in the season to evaluate survival. Maximum stem length, number of stems, and flower head production in grazed areas were equal to or greater than in the corresponding ungrazed areas at each of the three sites. The number of achenes per head was not significantly affected by grazing at Site 4, negatively affected at Site 6, and positively affected at Site 8. Trampling by cattle was a significant cause of damage only at Site 4 on the CPNA. Percent cover of herbaceous vegetation was similar over all sites and between grazed and ungrazed portions. Considering both statistical significance and observed trends, grazing seems to have been beneficial to L. congdonii reproduction on the Carrizo Plain and in the Kettleman Hills, but was neutral or detrimental on the Elkhorn Plain. •
1
CI; --- =cm I - --- -- C I___ C wO i C ..._0 C 00 - • -- - „ „ „ • ( a N I c i — e e i C en A I - ";" C a QE k — II - c (= i © c„ 1--- 1 - ■ i - i i - - a -- --
- = a ,,, - -cS • -1 S eg o O C c s; 11 c - - o „ i -a c i s —; a I --- O e-- 1- --- - • a k a c r - i 7 - - C s --- _ "--- c --- s c - -- © c " i c II II _. - e _ 11 e; 1 i i si s -
( -1 - ■- 11 - -S Ce = 1 C; i Y •••• Cgo c ; (3 c - 1 _ s e c C - • - 1 • g IS 1 e i C I 1 •; • ( c C II ;" • C i C - • c c e ; O o II 0 g c ----- •- - e c e ‘ -- --- i 11 11 -- O
I ---- I 1 = = i ci o -- a -- q 3 - -- i _ Q $o ■ E i- -- c•A g c II co a c -- k II - - g g ce g © ei a o" o ; o e e c c ,..-. - c •--- •- -- M e -ii ii - - C 1 i e ■ ‘ e s - -
- c3 ---- = - a 1 9 -c 7,- --" -; ti ' gi 1 ,....: 11m e 7; 7; „ 7-c I '7; 1_ c ;c O s ,,,o 3-9 1 C - a k II - -- k ci c Oes ----- 1 --- - c - e ; --■ ii 1 ii - C7 e •e e • o--- s - ( I 7 -1 W --1 e
e I = - 1 c C e = 7/ io _ -- 7 e; q _ e e -1 1 e C II ---- ; 7 g; - C ; a I ii "C --- ■,0 c c _ - _ -__ 11 e; ii i - e s C E c i c + i -- es - S ■I k 8? 9 „-- - i „ c, e CS ee c ‘ I I s - - -e - ■ e u S ; Q C5 c. c C U k " ___ = _ „ 1 - - c= -i c a e c 1 e ( 1 e ■I "-1 -- - c I --- e3 - 11 1 11 11 - - C 77 s -- c i -1 C
C = c i E 7 a c C = -"E 1 1 " = .6 - S ( c o _ o ca / o _ u = o - = u o - o - u C 1 C - i = i a c g = = s g Ea a- = u E C o ... 0: = , . = = E o 3 G 3 — o 1 - I- I = 1 o o -- 4 = 1 E = .2 = mi c E E E 3 5 -e g = c - g = o = c a 1 s E-. A c a = 17
II 1
a e Com a iso o em e ia co g o ii a i u es i g a e s u g a e a eas a Si e o e Ca i o ai Sa uis O is o Cou y Ca i o ia (see A e i A o e ac oca io Mea s a e s ow ± s a a e o Sam e si e is as i ica e a e o o eac co um u ess gi e i a e eses
SI E G A I G EGIME G A E U G A E I 1993 ( = 7 ( = 77 um e su i i g o ui i g 59/7 57/77 9 3 (75 (7 ca e 1 /7 /77 15 7 1 trampling ( 5 ( um e amage y ka ga oo 3/7 1 /77 9 a s ( 93 ( 3 u k ow 3/7 1 /77 3 11 7 sou ces (3 (13 Ma imum s em e g (cm 17 ± 1 19 1 ± 1 3 777 um e o s ems 5 ± 1 3 91 ± 33 5 5 o a owe ea o uc io 5 ± 1 17 3 9 ± 5 71 3 um e o ac e es e ea 1 1 ± 5 13 79 ± 3 5 15 ( = 1 7 (i = 1 9 . co g o ii e si y i e 3 • a sec ( a s/m e aceous co e ( 93 97 • •
• o es e 19
Table 9. Comparison of em e ia co g o ii attributes in grazed vs. ungrazed areas at Site 6 on the Elkhorn Plain San Luis Obispo County, California (see Appendix A for exact location). Means are shown ± standard error. Sample size is as indicated at the top of each column unless given in parentheses. values are from within-site precinct-by-grazing nonparametric analysis of variance.
SI E G A I G EGIME G A E U G A E P I 1993 ( = 1 ( = 7 um e su i i g o ui i g 1/1 / 7 97 (77 9 (7 ca e /1 / 7 1 1 trampling (3 ( i sec s /1 1/ 7 7 um e (1 9 (1 1 amage y ka ga oo 1/1 / 7 97 5 a s ( (3 u k ow /1 / 7 1 5 3 sou ces (1 9 ( Ma imum s em e g (cm 1 9 ± 97 15 5 ± 1 7 55 5 um e o s ems 5 ± 3 7 ± 3 3 3 o a owe ea 3 ± 7 3 5 ± 31 3 5 1 o uc io um e o ac e es e 1 9 ± 1 1 ± 7 3 59 ea (n = 3 (n = e aceous co e ( 99 9
Not tested.
a e 1 Com a iso o em e ia co g o ii a i u es i g a e s u g a e a eas a Si e i e Ke ema i s Ki gs Cou y Ca i o ia (see A e i A o e ac oca io Mea s a e s ow ± s a a e o Sam e si e is as i ica e a e o o eac co um u ess gi e i a e eses
SI E G A I G EGIME I 1993 G A E U G A E ( = 3 ( = ca e am i g 1/ 3 / um e ( 3 ( amage y u k ow sou ces 1/ 3 1/ ( 3 (3 Ma imum s em e g (cm 9 1 ± 1 ± 1 3 5 um e o s ems 9 ± 3 ± 1 7 o a owe ea o uc io 9 ± 1 31 33 9 ± 5 9 5 um e o ac e es e ea 3 5 ± 1 5 1 7 ± 1 37 9 9 ( = 1 ( = e aceous co e ( 99 9 -99
um e o occu e ces o sam e si es oo sma o e o m es 21 Survival, size, and reproduction of L. congdonii were investigated on giant kangaroo rat precincts and in interspaces at Sites 4 and 6 on the Carrizo and Elkhorn Plains, respectively. Plants growing on precincts were subject to significantly more damage from giant kangaroo rat activity (e.g., clipping, burial under loose dirt) than those in interspaces (Tables 11 and 12). However, survival was not affected by clipping because damaged plants generally persisted long enough to set seed. Plant success relative to occurrence on precincts differed between the two sites (site-by-precinct interaction; Table 13). Precincts did not significantly affect size of L. congdonii at the Carrizo Plain site (Table 11), but on the Elkhorn Plain individuals growing on precincts were larger, had more stems, and produced more heads than those between precincts (Table 12). Frequency of occurrence of L. congdonii on precincts was determined only at Site 4, where individuals were randomly distributed with respect to precincts (Table 11). Observations suggested that a similar distribution pattern occurred at Site 6 on the Elkhorn Plain.
The three-way interaction between sites, grazing, and precincts on the CPNA was significant for several L. congdonii variables (Table 11). Means for each cell are not presented, but inspection indicated that the highest mean values over all attributes were at the Elkhorn Plain, on precincts, in the grazed area. Low means were found under two combinations of conditions: (1) on the Elkhorn Plain, off precincts, in the ungrazed area, and (2) on the Carrizo Plain, on precincts, in the ungrazed area.
Habitat Characterization
Vegetation sampling was conducted on 19 transects on the CPNA and on 43 transects in other areas (Appendices B and C). Overall, vascular plant cover was high in 1993, and non-vegetated areas of bare soil or plant litter often were associated with kangaroo rat (Dipodomys spp.) activity. Cryptogams (e.g., moss, lichen, algae) were not often recorded on the transects. However, the low cover percentage of cryptogamic crust was to some extent an artifact of the sampling procedure, in which non-vascular plants were recorded only if the pointer did not hit living vascular plants or plant litter. On the sampled transects, the proportion of exotic plant species ranged from 16.2% in the Kreyenhagen Hills to 51.9% in the Panoche Hills and was 28.1% on the CPNA.
Cover and composition values were averaged over all transects that supported each listed species (Table 14). Transects in C. californicus populations had the greatest species richness and contained the lowest proportion of exotic plants. Opuntia basilaris var. treleasei occupied the most sparsely-vegetated sites. Listed species often were found in vegetation associations dominated by a particular plant species. Of the 14 transects in C. californicus populations, 8 were dominated by Vulpia myuros and 3 by V. microstachys. Eriastrum hooveri, Lembertia congdonii, and Gambelia sila often occurred together, and the majority of sites were dominated by V. myuros, Schismus spp., and/or Brornus madritensis ssp. rubens. Transects in Eremakhe panyi ssp. kernensis and Opuntia basilaris
Table 11. Comparison of em e ia co g o ii attributes on vs. off giant kangaroo rat precincts at Site 4 on the Carrizo Plain, San Luis Obispo County, California (see Appendix A for exact location). Means are shown ± standard error. Sample size is as indicated at the top of each column unless given in parentheses.
OCA IO E A I E O O O A ECI C ( = 7 ( = 5 um e su i i g o ui i g 5 /7 3 /5 1 93 (75 (7 ca e 5/7 7/5 9 3 am i g ( 9 (1 um e amage ka ga oo a s /7 7/5 3 13 y (3 1 (1 u k ow 3/7 7/5 1 7 sou ces ( (1 Ma imum s em e g (cm 17 55 ± 1 17 ± 1 7 9 3 um e o s ems 3 ± 1 ± 9 o a owe ea o uc io 3 7 ± 1 5 9 ± 11 1 3 535 um e o ac e es e ea 1 5 ± 31 1 31 ± 31 31 577 (n = 11 ( = 1 1 . co g o ii 71/1 9/1 um e o (5 7 ( 9 3 occu e ces 0.22 39 a om oi s /1 7 /1 ( 7 (5 3
Table 12. Comparison of Lembertia congdonii attributes on vs. off giant kangaroo rat precincts at Site 6 on the Elkhorn Plain, San Luis Obispo County, California (see Appendix A for exact location). Means are shown ± standard error. Sample size is as indicated at the top of each column unless given in parentheses.
OCA IO E A I E O O O ECI C ( = 55 ( = 73 um e su i i g o ui i g 7/55 5 /73 1 175 ( 5 5 (7 ca e 3/55 1/73 3 am i g (55 (1
um e i sec s /55 1/73 3 amage y (3 (1 ka ga oo 7/55 /73 5 a s ( 9 1 (3 1 u k ow /55 1/73 155 13 sou ces (73 (1 Ma imum s em e g (cm 15 ± 1 33 11 1 ± 9 35 3 um e o s ems 7 ± 7 ± 3 1 o a owe ea o uc io 9 35 ± 1 33 5 ± 7 3 5 um e o ac e es e ea 1 ± 1 15 ± 1 9 1 ( = 3 1 ( = 1
0 O ( O 4 ■ a 1 7 © c E- S C ei 1 -3 e co
E - e . ( c (" § E © © ©
( 1 I 1 g „ ;
,c, e a, c", a r- :x (kri i 1 x z O co Ei E I 1 I■1 ( I e ? - C c=: . / co co c
§ co Q+ § O & co © © ( I a c i I
( a g ©
; ; 7w
c § § g
; e 1 g C -- C•I o ©
e -- -■ co e © A - © S [1 1 co o c E- i I a e A A A C•1 e
e 5 1 A -o c c e - A
c a E = a a - e I " c 73 44 , 4.? 0 g O C — 7 o =• ; E 3 -3 E a ,... i i 1 a a 5
O 11 -
O O
M
o
O I § w --
a e easei o u a io s a so we e omi a e y . ma i e sis ss u e s. e e ce io o is a e was i o omys i ge s, w ic was ou i a i a s omi a e y a a ie y o a s ecies
a sec s a e C A we e oca e i wo a commu i ies ( o a 19 wi e ma o i y i U e So o a Su s u Sc u ( a e 15 Sig i ica y mo e e o ic a s ecies we e ese i o - a i e g ass a a i U e So o a Su s u Sc u a sec s o e Ca i o a E k o ai s i o i e sig i ica y i co e u e o me a a a ge um e o a s ecies ese i c u i g mo e e o ic s ecies a i e a e G a e a u g a e a eas a simi a ege a io co e u s ecies ic ess was owe a e o ics we e mo e omi e i g a e a eas O e a a sec s co e o e o ic a s accou e o e ma o i y o o a ascu a a co e a mo e a o e-qua e o a s ecies i e si y
e e o ic g asses Sc ismus a a icus, u ia myu os, a omus ma i e sis ss u e s a e e o ic o E o ium cicu a ium we e e a s ecies mos eque y e cou e e a omi a e (o co- omi a e o ma y a sec s o e C A ( a e 1 A oug ma y a i e a s ecies occu e o a sec s o y as e ia ca i omica a u ia ic os ac ys ac ie e omi a ce i a y o e C A sam es e a i e e e ia u c g asses oa secu a ss secu a (= . sca e a a a e (= S i a cemua we e ese o y o a sec s om e wes e ma gi o e C A a o y e o me co s i u e measu a e co e (a e agi g 5
ISCUSSIO
is o ic ag icu u a ac ices o e Ca i o ai i c u e y- a a mi g a i es ock g a i g ( uss ewis a oy a e oek e so a commu ica io s a i a mo i ica io esu i g om ese ac ices esuma y e o e e i a io o i i i ua co o ies o a s ecies ow is e as e a ge e Cau a us caqomicus was oug o a e ee e i a e om e Ca i o ai a e 195 ( wisse ma 195 oo e 197 ay o a a i a 19 u e s ecies was e isco e e y Mike os e i 19 ( oy a e oek e so a commu ica io A i io a o u a io s a e ee oca e o e C A e e y yea si ce 19 ( uss ewis e so a commu ica io oo e (197 e o e e occu e ce o . co g o ii o e Ca i o ai u i g e e io o i e si e use u o y a ew sma o u a io s we e o se e u i g e 19 s ( ay o 19 9 Co o ies o o s ecies may a e emai e u o ice i u cu i a e a eas
Cau a us ca omicus ow occu s ima i y i a eas o e C A a we e g a e u o a me w e eas e e si e co o ies o . co g o ii we e o e i 1993 i o g a e a e ious y cu i a e a eas o e C A ( ewis 1993 em e ia co g o ii a so as ee ou i e ious y cu i a e a eas o os i s ( ay o a uck 1993 o o u a io s o a ea i si es a we e a me ei e see s mus a e 7
O . • k ci o i n —1 -1- — c O 1 C3 E l i ri I C7 9 is-- • 7 C A 1 1 5 I c ■ --+ c g Q II a = 11 i 11 11 M 11 11 a 9
c o • • 1 g 7 7 91 g e ---- c _1 oo O oo — k ; -- 9 6 E . , ii 4 4 , . = t t 11 11 i o
a o c E -- 1 —• e -- - cc s 1 C e _ c= o 1 -Si 11 11 11
o — ■• p e c IQ oo s k k k t.'" 'tr C "I 71 tri i C: C e • ( -1 n —1 n — v. co — " M - = ∎ I 11 i 11 1♦ o 11 11 - c
e O --7 - s i c „ og r I a k e ei e ‘9 "" - _ 1 c= i ° 1 tr e M CA e C C • • e o I ■O i ii ii 11 11 " 11 11 u - C
- - - a ■ a --- -- S o — e — k e c --- --; cy co y — - ey A ago "-- - - -- - --• 44 (.... .i: , 4 C e -- C O O O O C O i 11 11 11 11 11 E - u
c C (1 c E I ■I ■0 C ■ 1■1 -I 11
a- a E c c a 4 ( 1 " g A i • 1— c 1 § - 1 e § g (i E o a (1 C . - o C MI (7 E ( = a o S a — a g C a J = U •- I 6 •
Table 16. Plant species most commonly encountered on 19 transects in endangered species populations on the Carrizo Plain Natural Area, San Luis Obispo County, California, in Spring 1993.
S ecies O igi A e age a so u e ese ce Co s a cy omi a ce co e ( 3 Sc ismus a a icus E 3 ± 155 1 1 11 E o ium cicu a ium E 7 7 ± 37 19 1 5 u ia myu os E 1 ± 1 1 a i s ua
o us ma i e sis E 175 ± 3 19 17 5 ss u e s as e ia ca i o ica 937 3 9 15 1 1 u ia mic os ac ys 1 ± 31 1 13 1 a auci o a o i oca um g aci e 9 ± 9 0 ec occuya e ici a a ± 1 19 1 1 e i i u ai u 11 ± 1 15 1 o us w a ge ia us 3 9 ± 1 5 1 1 A si ckia esse a a 3 7 ± 13 1 i o i u g aci e u 3 ± 1 1 1 a g aci e um o e u ai ma E 333 1 1 Ca a i ia ci ia a 3 ± 11 9 i a us i i o us 1 7 ± 13 E iogo um g aci i u m 1 3 ± 1
1 Nomenclature follows Hickman (1993). E = exotic, N = native. 3 Number of interceptions per species per 100 transect points. Number of belt transects on which the species occurred. 5 Number of transects on which the species was intercepted. Number of transects on which the species was dominant or co-dominant. 9 e sis e i e see a k o ece y is e se o e si e A ua s o a i e i o me s a e e o e a s a egy o oug esca e y s e i g e y seaso as see s ( o es e a 19 1 E e u i g e io s o o ima ge mi a io co i io s o a see s o suc s ecies ge mi a e a is s a egy o ec s e o u a io agai s u a o a e co i io s u i g e g owi g seaso o owi g e ge mi a io e io ( i i i 1993 See coa s o o C. californicus a L. congdonii equi e o o ge wea e i g e o e see o ma cy ca e oke (Ma e a e ickso 1993a Mo eo e Lembertia congdonii as imo ic see s a may i e i ge mi a io equi eme s ( ay o 19 9 us e e i g a see s om ge mi a i g i a gi e yea See a k s u ies wou e ea w e e u ie see is ese i is o ic oca io s a o o cu e y su o ese is e s ecies
e o se e a ia i i y i mo o ogy ecu i y a e o ogy amo g mo i o e o u a io s may e ue o a i a i e e ces oca a ia io s i wea e co i io s a /o ge e ic i e si y ( a y a e e ies 19 1 a commu i ies e e a io as ec soi s a is o ic a uses a y amo g e a eas (E e Cy e u u is e a a u e mo e eci i a io i e Sa oaqui a ey ec eases a o g o o -sou a wes -eas g a ie s (Ma o 19 e causes o a ia io amo g o u a io s a ei im ica io s o s ecies co se a io ca e e e mi e o y oug o g- e m emog a ic a mo o ogica s u ies i e si e sam i g o si e co i io s a a a ysis o ge e ic i e si y (Massey a W i so 19 a k a o si ge 1991 Owe a ose e e 199
o u a io es o ses o oug a ma ageme a so ca e e a ua e y co i ue mo i o i g e si y a o u a io es ima es om 1993 a e ese e as ase i e a ues o com a iso s wi o e yea s U o u a e y imi g a ec s e cou s ue o ea o i i i ua s oug e g owi g seaso Mo eo e e si ies ca a y g ea y e e wi i a si g e o u a io ue o e c um e is i u io o a s e si y es ima es may o i e a use u i e o a u a ce e wee yea s i ey a e com a e o e same e a sec a a e same e o ogica s age
O y g oss es ima es o a e a o u a io si e o e C A a e a ai a e om yea s ece i g e ece oug I ge e a o u a io s o o C. cali ornicus a L. congdonii a e ee i c easi g i si e i ece yea s ( uss ewis e so a commu ica io a oug ew see s o e a e s ecies ge mi a e i yea s o e ow- a e age ai a ( wisse ma 1 7 ay o 19 9 e ew o u a io s o L. congdonii o se e o e Ca i o ai i 19 i c u e a mos "se e a u e " i i i ua s eac ( ay o 19 9 w e eas i 1993 ma y o u a io s i c u e ousa s o a s ( ewis 1993
es i e e oss o some i i i ua o u a io s e Ca i o ai a u a A ea is a im o a si e o e o g- e m co se a io o o Caulanthus californicus a Lembertia congdonii. I is o e o e ee emai i g ce e s o co ce a io k ow o C. cali ornicus, a o g wi Sa a a a a Ca yo i Sa a a a a Cou y a e 30 Kreyenhagen Hills in Fresno County (Russ Lewis and Bruce Delgado, personal communications; California Department of Fish and Game 1991). In addition to the Carrizo Plain, primary areas for Lembertia congdonii are (1) the Panoche Hills of Fresno and San Benito Counties, (2) the Kettleman Hills of Fresno and Kings Counties, and (3) the Lost Hills area of Kern County (Russ Lewis, personal communication; Taylor 1989). The few populations of Eriastrum hooveri on the CPNA do not represent a significant portion of extant colonies. Major concentrations of this species occur in (1) the Cuyama Valley of Santa Barbara and San Luis Obispo Counties, (2) the Elk Hills-Buena Vista Hills-Lokern area of Kern County, (3) Lost Hills, and (4) the Kettleman Hills (Taylor and Davilla 1986, Lewis 1992).
The observed differences in rare plant performance between grazed and ungrazed areas on the CPNA may not be representative of grazing conditions in other years or at other sites. Studies of grazing effects on rare plants during 1993 were observational, not experimental, and did not control for intensity of use or possible differences in soil or microtopography between grazed and ungrazed portions of the sites. Grazing intensity varied even within portions of sites that were subject to cattle use, depending on forage availability and on the placement of livestock attractants such as water troughs and salt blocks (Ellen Cypher, personal observation). Therefore, these preliminary data should not be used as a basis for management decisions, but rather as indicators of the need for additional studies.
Giant kangaroo rats may promote growth of rare plants by creating favorable microhabitats or by dispersing seeds. Soils on precincts of another species of kangaroo rat, Dipodomys spectabilis, were higher in available nitrogen and lower in bulk density than soils of interspaces (Moorhead et al. 1988, Mun and Whitford 1990). Similar factors may have been responsible for the greater size and reproduction of L. congdonii on precincts on the Elkhorn Plain. However, if nitrogen is not the limiting factor at a particular site, occurrence on precincts may not affect plant size or reproduction. When moisture was limiting, plant cover did not differ between D. spectabilis precincts and interspaces (Moorhead et al. 1988). Perhaps limiting nutrients other than nitrogen regulated the size and reproduction of plants in Carrizo Plain populations of C. californicus and L. congdonii, which did not differ between precincts and interspaces.
Precinct age also may have affected plant response. Nitrogen accumulations in soils of recently-established precincts are likely to be appreciably lower than those in precincts used by generations of animals. Presence of giant kangaroo rats was verified at Sites 1, 2, and 6 prior to 1987, but Site 4 apparently was not inhabited at that time (Williams 1992). Although the relative ages of precincts at Sites 1 and 2 are unknown, precincts at the former typically were well-defined due to mounding, whereas those at the latter site were less distinct. Thus, degree of giant kangaroo rat activity may have influenced the growth response of C. californicus. 31 e is o o io a e occu e ce o C. ca4o icus o eci c s i o Ca i o ai o u a io s sugges s a gia ka ga oo a s may cac e see s o ui s o is s ecies Se ec i e a es i g o see s y gia ka ga oo a s ca ea o omi a ce o e e e a s ecies o eci c s (Sc i ma 199 Gia ka ga oo a s a a e y seek ou C. ca o icus, ecause s ems o is s ecies we e c i e wi equa eque cy o o eci c s a i i e s aces Co e se y c i i g o . co g o ii was imi e ima i y o a s occu i g o eci c s
a i e Ca i o ia g ass a s a e ee a e e om ese eme co i io s ue o i o uc io o e o ic a s i es ock g a i g cu i a io a i e (McC i ock 19 7 ea y 19 oug i e ac s wi o e cu i a io a o e g a i g o eg a e o ege a io a soi s; ege a io eg a a io i c u es c a ges i co e s uc u e a s ecies com osi io (G ai ge 199 e e ac e ec s o i es ock g a i g a e 19 7-199 oug o e ege a io o e C A a e u k ow ecause ase i e ege a io a a a e o a ai a e o com a iso u e mo e e ege a io a a ese e e ei a e o mea o e e ese a i e o e e i e C A o o e e i e a ge o a i a s o eac is e s ecies a sec sam i g was co uc e as a o mo e e e si e i es iga io s o is e s ecies a i a i e Sa oaqui a ey a a commu i y c a ac e is ics o e C A ese a a wi se e as a ase i e o e ea a e s o ege a io c a ge o e ime a ei e a io s i o o u a io si es o is e s ecies
Com a e o e s a e as a w o e e sam e a sec s a a ig e o o io o o - a i e a s E o ics com ise a o ima e y 11-1 o e a s ecies i Ca i o ia e e i g o e c i e ia use o e e mi e i c usio i e o a (McC i ock 19 7 a e 19 owe e e g ass a a s u a a i a s o e is e s ecies u e s u y a e ee im ac e mo e ea i y a o es e commu i ies y e o ic a s (Moo ey e a 19 o - a i e a s ike y ac ie e omi a ce o g e o e e ece oug (Wes e 19 1 Mos o e e o ic s ecies commo y e cou e e o a sec s ecame es a is e i Ca i o ia u i g e 19 ce u y ( ea y 19
Ac i e ma ageme suc as a i g a esc i e u i g cou i c ease e co e a is i u io o a i e a s o e C A a icu a y e e ia g asses a s u s I a i e s ecies e sis i e see a k a i g may o e ecessa y u ge mi a io o u wa e e o ics wou ee o e e e e ( a e a k a e e so 19 9 ega ess o e es o a io ec iques use e o ic a s a e o ike y o e e a ica e ( ea y 19 Sc i ma 199
Ma ageme o is e a s may ocus o e uci g com e i o s e a o s o a oge s a /o omo i g sym io ic s ecies suc as o i a o s see is e se s a myco i a u gi ( a y a e e ies 19 1 Messick 19 7 Mau e 199 O ima co i io s o eac s ecies a e u k ow e e mi a io o a o ia e ma ageme s a egies o e su e e o g- e m su i a o a e a s o o e Ca i o ai a 32 at other sites throughout their ranges, depends on site-specific experimental research (Bratton and White 1980, Massey and Whitson 1980, Davy and Jefferies 1981, Hodgkinson 1992).
RECOMMENDATIONS
Management decisions for the CPNA and other public lands should take into consideration the needs of listed plants and animals. However, directed research is necessary to make informed decisions. Aspects of rare plant biology and ecology that could be elucidated through further research include (1) site conditions affecting plant performance, (2) population responses to potential management techniques, (3) interactions with other plant and animal species in the community, (4) reproductive biology, and (5) genetic diversity within and between populations. For practical purposes, however, research topics and target species must be prioritized according to recovery needs.
Of the species included in this report, Cau a us ca o icus should receive highest priority for continued research because it has the most restricted distribution and the fewest individuals. The causes of differing plant success among C. ca i o icus sites should be investigated. Preliminary research may indicate whether site factors are limiting plant fecundity in the Kreyenhagen Hills or whether other factors should be considered, such as lack of pollinators or low genetic diversity. Data collection could be streamlined by focusing on reproductive plant characters, particularly survival to maturity, fruit production, and seed set. Research could be conducted either in the laboratory or in the field to evaluate the importance of site variables such as precipitation patterns, soil nutrient levels, and competition.
A secondary research need is to determine the impact of current livestock grazing practices on listed plants, particularly C. ca o icus and . co g o ii. Many of the known populations occur in present or former pastures, but resource personnel do not know whether or not to allow grazing to continue because empirical studies have not been conducted. Grazing studies should be conducted first in areas where tall, dense herbaceous vegetation may be outcompeting rare plants. The . co g o ii populations at Sites 5 and 8 met this criterion in 1993. At minimum, survival and fecundity of the listed plants should be compared between grazed and ungrazed treatments. Effective research requires a commitment from the administering agencies to manage grazing in specific study areas in order to meet rigorous statistical and experimental design criteria, which will require controlling livestock density, access to pastures, and seasons of use.
Finally, monitoring of rare plant populations should be continued through a full a ge of precipitation conditions, including years of drought, and normal and above- average rainfall. For each listed plant species, the number of individuals should be counted annually at designated sites to determine whether known populations of listed 33 plants are declining, increasing, or remaining stable. More intensive monitoring, including tracking survival and fecundity, will be necessary during and after any trials of burning, planting, or other management techniques in habitats occupied by endangered or threatened species.
Reintroduction of Caulanthus cegornicus, Lembertia congdonii, and Eriastrum hooveri on the CPNA is not recommended at this time because populations already are present. However, populations on the CPNA could provide seed sources for future reintroductions elsewhere if determined to be necessary for recovery. Any seed collections should be made according to guidelines established by the Center for Plant Conservation (1991) and only if appropriate permits have been issued by the U.S. Fish and Wildlife Service and the California Department of Fish and Game.
LITERATURE CITED
Bratton, S. P., and P. S. White. 1980. Rare plant management--after preservation what? Rhodora 82:49-75.
Brower, J. E., and J. H. Zar. 1984. Field and laboratory methods for general ecology. Second edition. William C. Brown Publishers, Dubuque, IA. 226pp.
Brown, J. H., and E. J. Heske. 1990. Control of a desert-grassland transition by a keystone rodent guild. Science 250:1705-1707.
California Department of Fish and Game. 1991. Annual report on the status of California state listed threatened and endangered animals and plants. California Department of Fish and Game, Sacramento, CA. 192pp.
California Native Plant Society. 1993. Field sampling protocol. Unpublished report. California Native Plant Society, Sacramento, CA. 5pp.
Center for Plant Conservation. 1991. Appendix: genetic sampling guidelines for conservation collections of endangered plants. Pages 225-238 in D. A. Falk and K. E. Holsinger, editors. Genetics and conservation of rare plants. Oxford University Press, New York, NY.
Davy, A. J., and R. L. Jefferies. 1981. Approaches to the monitoring of rare plant populations. Pages 219-232 in H. Synge, editor. The biological aspects of rare plant conservation. John Wiley and Sons, New York, NY.
Falk, D. A., and K. E. Holsinger, editors. 1991. Genetics and conservation of rare plants. Oxford University Press, New York, NY. 283pp. 3 G ai ge A 199 C a ac e i a io a assessme o ese i ica io ocesses ages 17-33 in G C a ma e i o ese i ie g ass a s ei io ogy a ma ageme Aca emic ess o o E g a
G i e 193 a i a e a io s o e gia ka ga oo a ou a o Mamma ogy 13 3 5-3
ea y 19 a ey g ass a ages 91-51 i M G a ou a Ma o e i o s e es ia ege a io o Ca i o ia 19 E i io Ca i o ia a i e a Socie y Sac ame o CA
eske E ow a Q Guo 1993 E ec s o ka ga oo a e c usio o ege a io s uc u e a a s ecies i e si y i e C i ua ua ese Oeco ogia 95 5 -5
ickma C e i o 1993 e e so ma ua ig e a s o Ca i o ia U i e si y o Ca i o ia ess e ke ey CA 1
o gki so K C 199 E eme s o g a i g s a egies o e e ia g ass ma ageme i a ge a s ages 77-9 in G C a ma e i o ese i ie g ass a s ei io ogy a ma ageme Aca emic ess o o E g a
o a 19 e imi a y esc i io s o e e es ia a u a commu i ies o Ca i o ia Ca i o ia e a me o is a Game Sac ame o CA 15
oo e 197 e ascu a a s o Sa uis O is o Cou y Ca i o ia U i e si y o Ca i o ia ess e ke ey CA 35
o es M M C u e a C Osmo 19 1 Mec a isms o oug esis a ce ages 15-3 in G a eg a As i a e i o s ysio ogy a ioc emis y o oug esis a ce i a s Aca emic ess Sy ey Aus a ia
ewis 199 Eriastrum hooveri ie i e o y U u is e e o o U S u eau o a Ma ageme ake s ie CA 11 + ma s
ewis 1993 Lembertia congdonii ie i e o y U u is e e o o U S u eau o a Ma ageme ake s ie CA O + ma s
Ma o 19 Ca i o ia c ima e i e a io o ege a io ages 11-7 in M G a ou a Ma o e i o s e es ia ege a io o Ca i o ia 19 E i io Ca i o ia a i e a Socie y Sac ame o CA
Massey a W i so 19 S ecies io ogy e key o a ese a io o o a 97-1 3 35 Mau e M 199 a ei o uc io a o e iew io i e si y a Co se a io 1 51- 1
Ma e S a e ickso 1993a emog a y a e o uc i e io ogy o Sa oaqui woo y ea s ( em e ia co g o ii: As e aceae e o o e Ca i o ia e a me o is a Game Sac ame o CA 5
Ma e S J., a A e ickso 1993 emog a y eco ogy a e o uc i e io ogy o Ca i o ia ewe owe (Ccu a us caW micus: assicaceae e o o e Ca i o ia e a me o is a Game Sac ame o CA 113
McC i ock E 19 7 e is aceme o a i e a s y e o ics ages 1 5-1 i S E ias e i o Co se a io a ma ageme o a e a e a ge e a s Ca i o ia a i e a Socie y Sac ame o CA
Messick C 19 7 esea c ee s o a e a co se a io i Ca i o ia ages 99-1 in S E ias e i o Co se a io a ma ageme o a e a e a ge e a s Ca i o ia a i e a Socie y Sac ame o CA
Moo ey A S am u g a J. A ake 19 e i asio s o a s a a ima s i o Ca i o ia ages 5 - 7 in A Moo ey a A ake e i o s Eco ogy o io ogica i asio s o o Ame ica a awaii S i ge - e ag ew Yo k Y
Moo ea F. M. is e a W G W i o 19 Co e o s i g a ua s o i oge - ic ka ga oo a mou s i a C i ua ua ese g ass a Ame ica Mi a a u a is 1 3- 7
Mu a W G W i o 199 ac o s a ec i g a ua a s assem ages o a e - ai e ka ga oo a mou s ou a o A i E i o me s 1 1 5-173
Owe W R., a ose e e 199 Mo i o i g a e e e ia a s ec iques o emog a ic s u ies a u a A eas ou a 1 3 -3
i i i 1993 e - e gi g ge mi a io o ese a ua s eyo e i s yea Ame ica a u a is 1 7 - 7
a e H. 19 e Ca i o ia o a ages 1 9-137 in M G a ou a Ma o e i o s e es ia ege a io o Ca i o ia 19 E i io Ca i o ia a i e a Socie y Sac ame o CA
Sc i ma M 199 omo io o e o ic wee es a is me y e a ge e gia ka ga oo a s ( i o omys i ge s i a Ca i o ia g ass a io i e si y a Co se a io I ess 3 ay o W 19 9 S a us su ey o Sa oaqui woo y- ea s (Lembertia congdonii). U S is a Wi i e Se ice Sac ame o CA 7 + A e ices
ay o W a E uck 1993 is i u io o Sa oaqui woo y- ea s (Lembertia congdonii) i e ici i y o os i s Ke Cou y Ca i o ia U u is e e o o os i s U i i y is ic os i s CA 17
ay o W a W a i a 19 S a us su ey o ee a s e emic o e Sa oaqui a ey a a ace a eas Ca i o ia U S is a Wi i e Se ice Sac ame o CA 131
wisse ma E C 195 A o a o e em o a ge a e eig o i g a o e Sa oaqui a ey Wasma ou a o io ogy 1 1 1-3
wisse ma E C 19 7 A o a o Ke Cou y Ca i o ia U i e si y o Sa a cisco Sa a cisco CA 395
U S e a me o e I e io is a Wi i e Se ice 199 E a ge e a ea e e wi i e a a s; e e mi a io o e a ge e o ea e e s a us o i e a s om e sou e Sa oaqui a ey e e a egis e 55(139 93 1- 937
a e a k A G a e e so 19 9 See a ks a e ma ageme a es o a io o a u a ege a io ages 3 9-3 i M A eck a ke a Sim so e i o s Eco ogy o soi see a ks Aca emic ess Sa iego CA
Wes e 19 1 Com osi io o a i e g ass a s i e Sa oaqui a ey Ca i o ia Ma o o 31- 1
Wi iams 199 Geog a ic is i u io a o u a io s a us o e gia ka ga oo a Dipodomys ingens ( o e ia e e omyi ae ages 3 1-3 in Wi iams S y e a A a o e i o s E a ge e a se si i e s ecies o e Sa oaqui a ey Ca i o ia ei io ogy ma ageme a co se a io Ca i o ia E e gy Commissio Sac ame o CA
Wi iams a K S Ki u 1991 Dipodomys ingens. Mamma ia S ecies 377 1-7
a 19 ios a is ica a a ysis Seco e i io e ice- a I c E g ewoo C i s 71 37 K OW E GEA E I I I UA S
Susa Ca e o a is Ca ie e esou ce A ea U S u eau o a Ma ageme 3 1 egasus i e ake s ie CA 933 - 37 ( 5 391-
uce e ga o o a is o is e esou ce A ea U S u eau o a Ma ageme ami o Cou o is e CA 95 3 ( 37- 1 3
G acie a i s aw ie io ogis S E a ge e S ecies eco e y a i g og am Sou C es e A e ue ake s ie CA 933 ( 5 3 -95
uss ewis Eco ogis Ca ie e esou ce A ea U S u eau o a Ma ageme 3 1 egasus i e ake s ie CA 933 - 37 ( 5 391-
Susa Ma e Assis a o esso e a me o io ogica Scie ces U i e si y o Ca i o ia Sa a a a a CA 931 ( 5 93- 11
o y e so ese e Ma age Ca i o ai a u a A ea O o 3 9 Ca i o ia a ey CA 93 53 ( 5 75- 3 3 au a Sc i ma Assis a o esso e a me o io ogy 1 111 o o S ee Ca i o ia S a e U i e si y o i ge CA 9133 ( 1 5-335
We y S e e s ie io ogis S E a ge e S ecies eco e y a i g og am 5 S ee ake s ie CA 933 1 ( 5 3 -5 7
oy a e oek 5 agu a S ee ake s ie CA 933 ( 5 3 -71 1 •
39
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7 5 Qua a g e u ic a su ey coo i a es ea me is e a sec (Si e ame s ecies co e ow s i a ge Sec io 1/ ese (Si e ES O COU Y K eye age i s 1S 15E E ca e E o eco A11 ( aca i os i s g a e (Si e 7 1S 15E 3 SE u g a e E o A3 eco A31
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S 1 E 1 W u g a e Caca K CC1 K CC Mo oc i e i ge 15S 1 E 3 E ? ii U SE U1 15S 13E 31 W ? u U SW U3
umey i s 15S 1 E SW u g a e 3 E iw eco A 1 ( a oc e i s ca e E o eco A g a e (Si e 1 KE COU Y Eas E k i s 3 S 3E E u g a e E o K1 E iso 3 S 3 E 19 W u g a e O S I SW S S 3 oke 9S E 1 SE s ee E ice K1 g a e 9S 3E 19 E u g a e ? E o K 9S 3E 9 W s ee E ke K11 g a e
Me e 11 W 3 SW u g a e ? O : K Mou o Ke 3 S 5E 1 SE u g a e E ke K Oi Ce e 9S E 1 E s ee O , K7 g a e
7.5' Quadrangle Public land survey coordinates Treatment Listed Transect (Site name) species code Township Range Section 1/4 present' (Site #)
Reward 29S 22E 32 SW ungrazed E ke K3 (Site 12)
Taft 31S 23E 10 SW ungrazed Gasi K5
West Elk Hills 30S 22E 2 NW ungrazed? E o Gasi K1 (Lokem) K2 (Site 13)
31S 23E 5 NW ungrazed? Gasi K4
KINGS COUNTY
La Cima 22S 17E 10 SW cattle eco KG1 (Kettleman Hills) grazed (Site 9)
Los Viejos 225 18E 26 SE ungrazed? E o KG3 (Kettleman Hills) 23S 18E 1 SW cattle E o Gasi LC6 grazed (Site 8)
23S 18E 2 NE cattle E o KG2 SE grazed eco Gasi LC2 (Site 8)
23S 18E 12 NW ungrazed eco Gasi LC1 eco Gasi LC3 E o eco LC4 Gasi E o Gasi LC5 (Site 8) SAN LUIS OBISPO COUNTY
Chimineas Ranch 31S 19E 11 NE ungrazed ii CP2
Cuyama lON 25W 9 SW ungrazed E o SLO
Painted Rock 32S 20E 8 NW cattle ii CP18 grazed 3
7 5 Qua a g e u ic a su ey coo i a es ea me is e a sec (Si e ame s ecies co e ow s i a ge Sec io 1/ ese (Si e a o ama i s 3 S E 1 W ca e ii Gasi C 1 (E k o ai g a e 3 S E E u g a e eco C 3 ii Gasi C ca e C 9 g a e C 1 (Si e We s a c 11 W W u g a e Caca ii C 13 (Ca i o ai 11 W 5 E u g a e Caca E ice C 15 W ii Caca ii C 17 11 W E u g a e Caca ii C 1 (Si e 1 11 W 9 W u g a e Caca ii C 19 U W 1 E u g a e Caca ii C 1
11 7W 11 E u g a e EliZO C U (Si e 3 11 7W 1 W u g a e E o C (Si e 3 U W 31 SE u g a e Caca E ice C 5 ii (Si e 3 S 1E 7 W ca e eco ii C 11 g a e (Si e 5 3 S 1E 35 SE ca e eco ii C g a e Gasi (Si e
3 S 1E 3 SW u g a e 5 eco ii C 7 Gasi (Si e SA A A A A COU Y o Mou ai 9 5W 11 SE u g a e Caca S U A E COU Y i ey 3S E SE g a e i i Gasi I 1 3S E 3 E g a e ? i i Gasi I E ie oo o es o A e i
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U •= 0 0 -■ oe -- e - a o u u 0s "I C 1 A S 1 ig S A e; - c i i o - -- -- i ? e ...._... i - 1- -- S c s o c- .40 oo C;51 o = c -e i cu - -- m - 0 O •-• e -I m 1 a 7i c c o o oo o & c O E-- a › › › 5 e e E e E e e ms m c c „s c a; i i i i i E -1= © — — ( I e m - oo m .0 1■1 1 - •••■1 e -1 i , 4 Si C o G G Q - Q - C A C c C - A 1 c 3 ( 51 _ 1 - u Qii C 1 CI i a &1 s c 1 a (3 1 c (
- - - " Q „ 4 „4. i . ,40 q 0,0 e O •-- e e - o ∎ — — - a a S a , . . 5 - 4 • ? 4 " t I t4 C: " I Y, s?. ° " I , : .4?, : . : e, a a a a a a c - U- - - U U U I U • " U U U U U U U .. U 9
(Cy 1 i = ce a 5
- Y A 47 1 1 E c " 1 i GO c CU --i
c C 2 g -; r :: 00 — — ■ =
i o $ oe ...., c o o o o o U 2 ,-„,,-, 5 e e k o o = cm -4
U C 1 ci C ssi © - a 1/ 1 u Q -i e; I c .. ? ; „E --- - v 1 15 (k -= a
-Eu = a - e Eg S ---k g E-- _ ao = › a u c cc c a ›- ca y - “C7 O — i e
C " c Ca = (1 k 5 a „ 5 a 111 • „7 Q44 c c II II II II II 11 II 9 00 ■I (u i a -fil 5 1 _ a ✓ i ” A 1---- E- i
50
APPENDIX D. Photographs of vegetation transects from the Carrizo Plain Natural Area, San Luis Obispo County, California. See Appendix B for exact transect locations. C 1 m C m
411.• CP3 0 m CP4 m
Pernoprn • C 5 m C m
111111111a aii
C 7 C (1 m C 9 m C 1 m
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- CP11 0 m CP12 0 m
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111•11111111Mme m _—____ CP13 0 m C 1 m
11111 1 111111111 C 15 m C 1 m C 17 m C 1 m C 19 Om
C 19 5 m