Dudleya cymosa subsp. pumila | |
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Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Eudicots |
Order: | Saxifragales |
Family: | Crassulaceae |
Genus: | Dudleya |
Species: | |
Subspecies: | D. c. subsp. pumila |
Trinomial name | |
Dudleya cymosa subsp. pumila (Rose) K.M.Nakai | |
Synonyms | |
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Dudleya cymosasubsp. pumila, most commonly known as the low canyon dudleya, [2] chalky canyon dudleya [3] or California live-forever, [1] is a species of perennial succulent plant. It has diamond to spoon shaped leaves, sometimes coated with a fine white powder, and in May through July, bright red, orange or yellow flowers adorn the short inflorescence. [4] A leaf succulent primarily found growing in rocky cliffs and slopes, it is endemic to California, and grows in the Transverse Ranges and South Coast Ranges, with some outlying populations. A variable plant, in some localities it is difficult to distinguish from other plants in the genus. [5]
The plant has diamond to spoon shaped leaves, with a height of 5 to 15 cm, and reddish-orange or yellow flowers. [6]
The stem is typically 1 to 2 cm wide, crowned by an apical rosette. The rosettes are often solitary, although some plants may have branching stems with 2-5 rosettes. Each rosette is 4 to 10 cm wide, with 10 to 25 leaves. The leaves are green, and some are glaucous or even farinose, although few are green in times of drought. The leaves are shaped rhombic-oblanceolate to spatulate, with the leaf margins (edges) folding up at the widest point, and the tip shaped generally short-acuminate to mucronate, more or less recurved. The leaves are 1.5 to 5 cm long, and 10 to 30 mm wide. [5] [4]
The inflorescence is more or less symmetrical on the radial axis. The peduncle is 5 to 30 cm tall, and 3 to 10 mm wide, and erect. There are 5 to 20 bracts on the floral shoots, with the lower bracts more or less not plump. The internodes are greater than 5 mm distant from each other. There are generally 3 first degree branches on the inflorescence, and the terminal branches are 1 to 3 cm long, with 3 to 6 flowers. The flowers have bright yellow to red petals. [4]
The majority of the plants in this species are diploid (n = 17), meaning they have two sets of chromosomes. Some plants found at a lower altitude may resemble this species, but have a chromosome count of (n = 34), making them tetraploid, or with four sets of chromosomes. As the species Dudleya lanceolata is tetraploid [7] and often sympatric with other species such as D. c. subsp. pumila, these plants are instead placed within lanceolata. [8]
The holotype was collected by H. M. Hall in the San Bernardino Mountains, with the taxon originally described as Dudleya pumila by Nathaniel Lord Britton and Joseph Nelson Rose during their revision of North American Crassulaceae. In 1902, Hermann E. Hasse collected a plant growing on the rocky banks of San Gabriel Canyon at 600 to 700 m in altitude. Hasse's plant was then described by Britton and Rose as Dudleya minor. [9] In 1957, botanist Reid Moran combined Dudleya minor under Dudleya cymosa, forming Dudleya cymosa subsp. minor. [10] The majority of Dudleya cymosa plants in Southern California were thereafter assigned to this subspecies. [8]
Research by botanist Kei M. Nakai showed that the type specimen of Dudleya minor, along with plants at the type locality, in the lower altitudes of San Gabriel Canyon, were in fact representative of Dudleya lanceolata, on the basis of morphology and chromosome count. This meant that the Dudleya pumila specimen by Hall was the accurate representative of Dudleya cymosa plants in the range. Nakai then combined Dudleya pumila into Dudleya cymosa subsp. pumila. The species Dudleya lanceolata and Dudleya cymosa can be separated on the basis that lanceolata is tetraploid and occurs at lower elevations, while cymosa is diploid and occurs at higher elevations. [8]
A phylogenetic analysis in 2013 did not provide any major revelations for the position of this subspecies, placing it as unresolved but emerging from a large polytomy at the base of the generic tree. Others members of the species Dudleya cymosa were better resolved, such as Dudleya cymosa subsp. paniculata and Dudleya cymosa subsp. cymosa, which are closely related with Dudleya abramsii subsp. setchellii. [11]
Hybrids with Dudleya lanceolata and Dudleya caespitosa are suspected. [4]
This species is endemic to the state of California. It has been reported from the Santa Lucia Range in Monterey County south to the San Gabriel and San Bernardino Mountains of southern California. [8] It is also found on Santiago Peak in the Santa Ana Mountains of western Riverside and Orange County. [3] It occurs on rocky cliffs and slopes. [4] It is found 100 to 1800 meters from Santa Barbara County north, and from 600 to 2600 meters from Ventura County south. [8]
The Sonoran blue (Philotes sonorensis) butterfly uses D. cymosa subsp. pumila [Note 1] [12] as a larval foodplant, with the butterfly closely following this species' range from Monterey County through the Transverse Ranges. [13] The butterfly lays the eggs on the upper parts of the leaves near the margin, and on the small leaves near the apical meristem at the center of the rosette. Plants with the eggs are usually in exposed areas, rather than partial shade. Once the larvae emerge from the eggs, they burrow into the Dudleya leaf in close proximity to their egg, feeding head down and burrowing deep within the leaf. Others may crawl over the leaf surface. The larvae may be spotted due to the disturbed leaves and frass left behind. [13]
The relationship of this butterfly with Dudleya potentially has taxonomic implications for the plants. Botanist Reid Moran described three subgenera of Dudleya; Stylophyllum, Dudleya, and Hasseanthus. The subgenus Dudleya consists of broad-leaved rosette-forming plants with connate petals adapted to hummingbird pollinators. Subgenera Stylophyllum and Hasseanthus have broad, open flowers, with Stylophyllum consisting of evergreen plants with pencil-shaped leaves, and Hasseanthus consisting of deciduous, geophytic plants. [13] This species is a member of the subgenus Dudleya. [14]
Entomologist Oakley Shields noted an emerging pattern when examining the Dudleya foodplants of Philotes sonorensis, as most of the foodplants are evolutionarily intermediate species within the genus. This species in particular has chromosomal resemblance to the Hasseanthus clade, which appears to place it as a primitive member of the subgenus Dudleya. [13] The subgenus Dudleya itself may also have evolved from the subgenus Stylophyllum , as indicated by the relationships between Dudleya stolonifera (a primitive member of the Dudleya subgenus) and Dudleya edulis (an advanced member of the Stylophyllum subgenus), [15] [16] both of which are used by the butterfly. In contrast, other species such as Dudleya pulverulenta, regarded as the most advanced member of the genus, and Dudleya blochmaniae, a primitive Hasseanthus plant, are not utilized by the butterfly. [13]
In spite of the perceived patterns between the butterfly's foodplants and the taxonomy of the genus Dudleya, recent phylogenetic analysis has cast doubt on the artificial divisions of Dudleya. [11] The Hasseanthus species may not be "primitive" and in fact may represent a derived subgroup with plesiomorphic features, [14] Reid Moran and Shields had noted the taxonomic difficulties and the failure of the subgenus Dudleya to be easily differentiated. [13] [15]
Hummingbirds feed on its nectar. [2]
Dudleya, commonly known as liveforevers is a genus of rosette-forming succulent plants in the stonecrop family, Crassulaceae, consisting of about 68 taxa in southwestern North America and Guadalupe Island. The species come in many forms, some large and evergreen, others geophytic and deciduous. Yet, despite their dramatic variations in appearance, most species readily hybridize. The flowers of Dudleya have parts numbered in fives, with the petals arranged in tubular, star-shaped, and bell-shaped forms and, when fruiting, are filled with tiny, ovoid-crescent-shaped seeds.
Dudleya abramsiisubsp. setchellii, known by common name as the Santa Clara Valley dudleya or Santa Clara Valley liveforever, is a member of the Dudleya genus of succulent perennials, members of the family Crassulaceae. The Santa Clara Valley dudleya, endemic to the Santa Clara Valley region in the southern San Francisco Bay Area, was listed on 3 February 1995, as an endangered species. It is considered to be a subspecies of Dudleya abramsii, but its taxonomic status is still unclear. Its closest relative is Dudleya cymosa subsp. paniculata, which is a morphologically similar sister taxon.
Dudleya cymosa is a species complex of evergreen and deciduous succulent plants in the family Crassulaceae known by the common name: canyon liveforever. It is a loosely defined polyphyletic species with a diverse number of subspecies, varying highly in morphology, distribution, and habitat.
Dudleya caespitosa is a succulent plant known by several common names, including sea lettuce, sand lettuce, and coast dudleya. It is endemic to California, where it grows along the coastline in the southern half of the state. Taxonomically, this species is a highly variable complex of polymorphic and polyploid plants, closely related to numerous neighboring species such as Dudleya farinosa, Dudleya greenei and Dudleya palmeri. It is delimited from neighboring species on an arbitrary basis of distribution and chromosome number, and may not be immediately separable from the other species it approaches.
Dudleya abramsiisubsp.calcicola is a succulent plant known by the common name limestone liveforever, or limestone dudleya. It is endemic to California, where it is a rare resident of limestone outcroppings and rocky slopes in the southern Sierra Nevada and nearby mountains and foothills. It was formerly regarded as Dudleya calcicola.
Dudleya farinosa is a species of succulent plant in the family Crassulaceae known by several common names, including bluff lettuce, powdery liveforever, and powdery dudleya. A coastal plant of northern California and southern Oregon, it is typically found on ocean bluffs just directly above the reach of the waves, and sometimes inland. Its appearance is characterized by lotus-like rosettes of beveled leaves, and in summer the plant erects a tall pink to red peduncle densely covered in bracts, topped with branches of pale yellow flowers. The green or white rosettes of this plant can be seen covering stretches of rocky coast and nearby islets.
Dudleya lanceolata is a succulent plant known by the common name lanceleaf liveforever or lance-leaved dudleya. It is an extremely variable and widely ranging species that occurs from Monterey County and Kern County in California south through Ensenada in Baja California. It is characterized by green to purple lanceolate leaves, red, orange, or less commonly yellow petals, and is typically tetraploid. Despite its diversity, it is quite stable as a species, but hybrids may be discovered with other species of Dudleya, which can make it difficult to discern in areas where numerous species converge.
Dudleya pulverulenta is a species of perennial succulent plant known by the common names chalk lettuce, chalk dudleya, and chalk liveforever. It is one of the largest Dudleya, with a silvery, waxy rosette that may greatly contrast with its habitat. It is also regarded as one of the most distinctive members of the Dudleya, with one of the most specialized inflorescences in the genus, adapted to hummingbird pollination through its red pendent flowers, the longest corolla, and the highest nectar output. Dudleya pulverulenta has the largest range of all Dudleya, over 1,000 kilometres (620 mi), being found from southern Monterey County in California to the Sierra de San Borja in southern Baja California. It is closely related to Dudleya arizonica, a smaller desert species that tends to lack the specialized floral traits, and Dudleya anthonyi, which differs in a few morphological traits and is restricted to the San Quintín Volcanic Field.
Dudleya stolonifera is a succulent plant known by the common name Laguna Beach liveforever or Laguna Beach dudleya. This is a rare plant which is endemic to the coastline of Orange County, California. It is known from only about six populations in the vicinity of Laguna Beach, totaling about 30,000 individuals. It is federally listed as a threatened species of the United States.
Dudleya traskiae is a rare succulent plant known by the common name Santa Barbara Island liveforever. This Dudleya is endemic to Santa Barbara Island, one of the Channel Islands of California, where it grows on rocky bluffs. The plant has a basal rosette of flat, spade-shaped fleshy leaves up to 15 centimeters long, which are pale green to yellowish. It erects tall stems bearing dense, rounded inflorescences of many bright yellow flowers.
Dudleya greenei is a perennial species of succulent plant known by the common names Greene's liveforever, or Greene's dudleya. It is endemic to the Channel Islands of California, where it grows along the cliffs of four of the eight islands. It is a highly variable plant, presenting with multiple forms and varying levels of ploidy. Taxonomically, this species is an insular segregate of Dudleya caespitosa, and was placed as a stopgap taxon by Reid Moran in his 1951 thesis on the genus. It is characterized by white or green leaf rosettes, loomed over by inflorescences bearing pale yellow to white flowers. It is a member of the subgenus Dudleya, as it cannot be propagated from leaf cuttings, does not grow from a corm, and has tight petals.
Dudleya variegata is a deciduous succulent plant in the family Crassulaceae known by the common names variegated liveforever, variegated dudleya or rarely San Diego Hasseanthus. A cryptic plant that survives part of the year dormant underground from starch reserves in a corm, after sufficient rainfall, leaves will emerge, soon giving way to small inflorescences with yellow star-shaped flowers. It is native to Baja California and adjacent San Diego County in California, where it grows in several habitat types, including chaparral and vernal pools.
Dudleya attenuata is a species of perennial succulent plant known by the common name taper-tip liveforever, native to Baja California and a small portion of California. It is a rosette-forming leaf succulent which has narrow pencil-shaped leaves that can often be found covered in a white epicuticular wax. The thin, sprawling stems branch to form the clusters of rosettes, with plants creating a "clump" up to 40 cm wide. The small flowers are white or yellow, with 5 spreading petals. It is a diverse, variable species that extends from the southernmost coast of San Diego County to an area slightly north of the Vizcaino Desert, hybridizing with many other species of Dudleya in its range. Some plants with white or pinkish flowers were referred to as Orcutt's liveforever, referring to a former subspecies split on the basis of the flower color.
Dudleya ingens is a species of perennial succulent plant in the family Crassulaceae commonly known as the rock liveforever or Baja liveforever. A relatively large member of the genus Dudleya, this species has long green succulent leaves, and in April to June is characterized by pale yellow to white pink-tinged flowers topping tall, reddish inflorescences. It has a stem clothed densely with old, leathery leaves, and the inflorescence may be nodding, with the floral branches bearing the flowers tending to unfurl like the fronds of a fern. It is similar in appearance to Dudleya brittonii, but differs in range and chromosome number. This species is endemic to the state of Baja California in Mexico, being found from Santo Tomás to the southern coast of the state.
Dudleya anomala is a rare species of succulent plant in the family Crassulaceae commonly known as the Todos Santos liveforever. With a dense, cushion-forming habit, this leaf succulent is characterized by elongated stems, slightly sticky leaves, and bell-shaped flowers with white, spreading petals. This species is native to Baja California, Mexico, and is found primarily on islands and one coastal locality.
Dudleya formosa, known by the common name La Misión liveforever, is a species of perennial succulent plant endemic to the Guadalupe Valley in Baja California. It is characterized by bright green leaves, red floral stems, and pink flowers.
Dudleya saxosa subsp. collomiae, known by the common name Gila County liveforever, is a subspecies of perennial succulent plant within the genus Dudleya native to central Arizona. It is characterized by showy bright-yellow flowers on an upright inflorescence colored pink, red or orange. The leaves are green or covered in a white, powdery wax. This species is found growing in rocky slopes, canyons, and crevices, and often on Sonoran Desert sky islands.
Dudleya cymosa subsp. paniculata, known by the common name Diablo Range dudleya, is a species of perennial succulent plant in the family Crassulaceae native to the Inner South Coast Range of California. It is characterized by pale yellowish flowers, oblong to oblanceolate leaves and a growth habit not limited to a single substrate. It is closely related to Dudleya abramsii subsp. setchellii.
Dudleya saxosasubsp. aloides is a species of perennial succulent plant in the family Crassulaceae known by the common names desert dudleya or desert savior. It is a rosette-forming species widely distributed throughout the Peninsular Ranges and desert mountains of California in the United States. It is characterized by bright-yellow or greenish-yellow flowers, and can be found in shaded crevices and slopes. Plants in western half of the range may grade into Dudleya lanceolata.
Dudleya cymosasubsp. cymosa is a species of succulent perennial plant in the family Crassulaceae native to California. It is the autonymous subspecies for Dudleya cymosa, and is known by the common name canyon liveforever. It is native to the California Coast Ranges, the Sierra Nevada and the Santa Monica Mountains. It is characterized by bright-yellow, orange or red flowers and broad, wide leaves. This plant is commonly found growing on rocky outcrops, talus slopes, and in shaded canyons.
On the basis of 18 or more well-scattered collections, it is tetraploid; it seems best defined partly on that basis.
Dudleya cymosa is a highly variable taxon that occurs throughout the foothills of California, and has been treated as nine different subspecies (McCabe 2012). Many of the relationships among subspecies remain unresolved with D. c. subsp. agourensis K. M. Nakai, D. c. subsp. ovatifolia (Britton) Moran, D. c. subsp. crebrifolia K. M. Nakai & Verity, D. c. subsp. pumila (Rose) K. M. Nakai, and D. c. subsp. marcescens Moran stemming from a large polytomy at the base of the tree. However, two of the D. cymosa subspecies, D. c. subsp. cymosa and D. c. subsp. paniculata, occur in the Virens clade as sister to D. abramsii subsp. setchellii, a federally listed endangered species from Santa Clara County, California.