The Double Cone Quarterly
Window to the Wilderness
Spring Equinox 1999 || Volume II, Number 1

Unique and Noteworthy Plants
of the Santa Lucia Mountains

Part One:
The Endemic Bedstraws of the Santa Lucia Mountains

By David Rogers

The purpose of this series is to help make familiar to aficionados of the Santa Lucia Mountains the many plants of this region that are noteworthy because they are (1) endemic (limited to) these mountains, (2) they are montane (mountain) plants that are disjunct (separated) from the nearest populations in other mountainous areas of California (sometimes radically so), (3) they are plants that reach their most southern natural distribution in these mountains; and (4) plants that are at their most northern natural distribution in these mountains. I will also discuss nearly endemic plants, i. e., species or lesser taxa that are almost entirely restricted to the Santa Lucia Mountains, but are also represented in nearby mountain ranges, such as the Santa Cruz, Gabilan, La Panza and Santa Ynez Mountains.

As this article pertains to a few of the endemic plants of this mountain range, I will provide a brief overview as to why a relatively large number of species are confined to this region. The Santa Lucia Mountains are centrally located on the coast of the California Floristic Province, which is one of the most noteworthy geo-botanical regions on earth, not only because of its high degree of endemism (47.7% according to Raven & Axelrod, 1978), but also because of the large number and diversity of its species. Major factors contributing to this rich assemblage of plants include a mediterranean climate, great variations in topography, and a wide varitey of soil types. In general, the CFP (aka Cismontane California) consists of the portion of the state west of the axis of the Cascade, Sierra Nevada, Transverse and Peninsular ranges, and includes southwestern Oregon and northwestern Baja California. The province also includes the area around Lake Tahoe and the islands off the coast of southern California and northern Baja California, and excludes the Modoc Plateau of northeastern California and the deserts of the southeastern portion of the state. Within the CFP there are a number of smaller regions that stand out due to their high number of even more narrowly of endemic plants, one of which is the Santa Lucia Mountains of Monterey and northwestern San Luis Obispo County.

The Endemic Bedstraws of the Santa Lucia Mountains

Three rare taxa of the bedstraw genus Galium are endemic to the Santa Lucia Mountains: two highly distinct species and a lesser taxon of the Galium californicum complex. While the common name for this genus refers to the use of some species as bedding materials (many are mat-forming plants with short, slender and yielding stems), the botanical name is derived from the Greek word for milk (gala), for certain species were used as curdling agents. Galium is a widely distributed (but primarily temperate) genus comprised of about 400 species, which is represented in the State of California by 39 native species plus 29 lesser taxa, 56% of which (38 taxa) are endemic to the California Floristic Province (these figures do not include taxa that are limited to the portions of the CFP in southwestern Oregon and northwestern Baja California). The genus is one of the approximately 500 genera of Rubiaceae (the Madder Family), a large ( 6,000 species) and primarily tropical family of trees, shrubs and herbs. Well-known plants of this family include Coffea (coffee), Cinchona (from which quinine is derived), and a number of common ornamentals, such as Gardenia, Pentas, Houstonia, Bouvardia and Hedyotis.

Galium clementis Eastwood.
Santa Lucia Bedstraw.

Small and densely leafy perennial herbs with numerous slender and generally erect stems ranging from about 8 to 13 cm. (3-5") tall. As the stems rise from a system of slender rhizomes, the plants form mats which can cover many square feet. The light green and finely hispid-pubescent leaves are produced in whorls of four (or occasionally six) at the nodes, and the blades, which are about 2 to 7 mm. long, are narrowly ovate-lanceolate to elliptic-oblong, but appear to be linear due to strongly revolute margins. The flowers are very small and produced in the axils of the upper leaves, the staminate in small cymes and the pistillate singular, and the corollas are four lobed, pale-yellow, and about 1 mm. wide. The fruits are two-lobed, fleshy, and about 1 to 2 mm. wide.

Galium clementis is scattered in suitable habitats above about 3000 feet in the Santa Lucia Mountains of Monterey County, and all of the known populations occur within the boundaries of the Monterey Division of Los Padres National Forest. To the best of my knowledge, this species has been collected at only seven sites, six of which are in the Ventana Wilderness, and the other in the Silver Peak Wilderness. The type specimen was collected by Mary Strong Clemens while on an expedition between Tassajara Hot Springs and The Indians, via the Santa Lucia Trail, in early October of 1921. Twelve years later Alice Eastwood, of the California Academy of Sciences in San Francisco, recognized that the specimen represented a very distinct species hitherto unknown to science, which she named in honor of its collector (Eastwood 1933). G. clementis is probably a relictual (ancient) species that is to some degree ancestral to the G. californicum complex.

Galium hardhamae Dempster.
Clare Hardham's Bedstraw.

Low, matted and sparsely hispid-pubescent perennial herbs with ascending or weakly climbing stems less than 30 cm. long. The fleshy leaves are produced in remote whorls of six at the nodes, and the blades, which are about one to three mm. long, are lanceolate to ovate, acute, and tipped with a stout hair. The colorless petioles are moisture-sensitive, thus the leaves spread horizontally when wet but become downturned when dry. The flowers are produced in small cymes in the axils of the upper leaves, and the rotate corollas are yellow or green, and about 2 mm. wide. The fruits are fleshy, two-lobed, and about 1 to 2 mm. wide.

Galium hardhamae is restricted to serpentine outcrops from the Silver Peak Wilderness and southward into northwestern San Luis Obispo County, usually in or near groves of Sargent Cypress (Cupressus sargentii), and at elevations of about 2000 to 3000 ft. The species is named for Clare Hardham, who discovered it in the late 1950s (Dempster 1962). Mrs. Hardham, of Paso Robles (San Luis Obispo County), has for many years been a tireless student of the native flora of central California, and in the process she has made major contributions to the understanding of the plants of this region, including the discovery of a number of previously unknown taxa. G. hardhamae is unique among the fleshy-fruited Galium species of the California Floristic Province in having six leaves per node instead of four. Its closest relative appears to be the other endemic Galium of the Santa Lucia Mountains, G. clementis, which occasionally has six leaves per node. Another dissimilarity between these two species and other fleshy-fruited Galiums are the petiolate leaves. It is possible that the two species shared a common ancestor, and diverged by adaptation to different soil types (soils derived from granitic and metamorphic rock for G. clementis, and soils derived from serpentine rock for G. hardhamae).

Galium californicum Hooker & Arnott subspecies luciense Dempster & Stebbins.
Coast Ridge Bedstraw.

Small grayish-pubescent perennial herbs with numerous stems ranging from about 5 to 16 cm. (2-6.5") long. As the stems rise from extensive underground rhizomes, the plants tend to form mats that can spread over an area of many square feet. The slightly fleshy leaves, which are produced in whorls of four at the nodes, are elliptical to slightly obovate, about 4 to 6 (-10) mm. long, and rather abruptly taper to a petiolar base. Staminate flowers are produced in few-flowered terminal and axillary clusters, while pistillate flowers are usually solitary in the axils of the upper leaves. The corollas are four-lobed, rotate, pale yellow or greenish, and only a few mm. wide, while the dark and fleshy fruits are two-lobed, densely pubescent, and only slightly wider than the corollas.

G. californicum luciense is restricted to the intermediate and higher elevations of the Coast Ridge, where it occurs in semi-shady areas in pine and oak forests. It represents one of the seven subspecies of the G. californicum complex, and was probably derived through genetic interchange between G. clementis and G. californicum flaccidum. The small leaves with petiolar bases are the most pronounced morphological characteristic of G. c luciense that is intermediate between G. clementis and G. c. flaccidum. At lower elevations it is often difficult to distinguish from small leafed forms of G. c. flaccidum, which, in the Santa Lucia Mountains, is a highly variable entity.

The morphological characteristics of the local representatives of another subspecies of the G. californicum complex, G. c. maritimum Dempster and Stebbins, are also noteworthy. This taxon, which is limited to coastal bluffs and canyons from Monterey County to Santa Barbara County, is most noticeably different from the other subspecies by its semi-woody stems, which climb or clamber on other plants. This characteristic is most pronounced in the plants along the Big Sur coast.

For systematically inclined readers I have prepared the following key to the perennial bedstraws of the Santa Lucia Mountains:

1a.  Leaves produced in whorls of five or six, or at least the lower 
     and middle leaves produced in whorls of 5 or 6:
   2a.  Leaves produced in whorls of four to six.  Corollas three-lobed...  
                                                       G. trifidum pacificum.
   2b.  Leaves produced in whorls of six.  Corollas four-lobed:
      3a.  Leaves 6 to 38 mm. long.  Often found in redwood forests...
                                                                G. triflorum.
      3b.  Leaves 1 to 3 mm. long.  Restricted to serpentine soil, usually
           in cypress forests...                                G. hardhamae.
1b.  Leaves produced in whorls of four (or occasionally six in G. clementis):
   4a.  Fruits densely bristly...                           G. angustifolium.
   4b.  Fruits glabrous or pubescent, but not bristly:
      5a.  Leaves awl-like, narrowly lance-linear, and sharp to the touch. 
           Dry areas towards the Salinas Valley...              G. andrewsii.
      5b.  Leaves not awl-like, not narrowly linear, or if so, then not sharp
           to the touch:
          6a.  Leaves linear to linear-oblong, or appearing to be linear due 
               to strongly revolute margins:
              7a.  Short mat forming plants with herbaceous stems less than 
                   15 cm. tall.  Leaves 2 to 7 mm. long and appearing to be 
                   linear because of strongly revolute margins...  
                                                                G. clementis.
              7b.  Subshrubs with semi-woody stems up to 200 cm. long, which 
                   tend to climb on other plants.  Leaves 2 to 18 mm. long and 
                   with flat or only slightly revolute margins...  
                                                          G. porrigens tenue.
          6b.  Leaves not linear or appearing to be linear:
               8a.  Plants with long, semi-woody and usually climbing stems:
                  9a.  Stems and leaves pubescent.  Restricted to coastal bluffs
                       and canyons...              G. californicum maritimum.
                  9b.  Stems and leaves glabrous or nearly so.  Widespread...  
                                                      G. porrigens porrigens.
               8b.  Plants with short, non-woody or only slightly woody, and 
                    usually tufted stems:
                  10a.  Leaves mostly 4 to 6 mm. long, more or less fleshy, and 
                        abruptly tapering to a petioled base.  Stems 5 to 16 
                        cm. long...                 G. Californicum luciense.
                  10b.  Leaves 4 to 25 mm. long, not fleshy, and without a 
                        petioled base.  Stems 7 to 32 cm. long:
                        11a.  Leaves with more or less sparse, scattered hairs, 
                              and rounded to abruptly acute at the tip...
                                                 G. Californicum californicum.
                        11b.  Leaves with dense, soft, fine hairs, or sometimes
                              nearly glabrous, the tip rounded to obtuse...
                                                    G. Californicum flaccidum.


DEMPSTER, LAURAMAY. Rubiaceae in The Jepson Manual, Higher Plants of California, James Hickman, editor. University of California Press. 1993. Rubiaceae, vol. 4, part 2, of Willis Linn Jepson's The Flora of California. Jepson Herbarium, University of California, Berkeley. 1979. A New Species of Galium in California. Madrono 16: 166-168. 1962.

DEMPSTER, LAURAMAY, and G. LEDYARD STEBBINS. A Cytotaxonomic Revision of the Fleshy-Fruited Galium Species of the Californias and Southern Oregon (Rubiaceae). University of California Publications in Botany 46: 1-57. 1968. The Fleshy-Fruited Galium Species of California. 1. Cytological Findings and Some Taxonomic Conclusions. Madrono 18: 105-113. 1965.

EASTWOOD, ALICE. Two New Species of Western Galium. Leaflets of Western Botany 1 (6): 55-56. 1933.

RAVEN, PETER, and DANIEL AXELROD. Origin and Relationships of the California Flora. University of California Publications in Botany 72. 1977.

YADON, VERN. Two annotated lists of plants observed by Mr. Yadon while on botanical forays in the Santa Lucia Mountains in 1979. Mr. Yadon also discovered another previously unknown locality for one of the rare Galium species in July of 1967, as evidenced by a specimen on file at the California Academy of Sciences.

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