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Dudleya

Dudleya, commonly known as liveforevers, is a genus of rosette-forming succulent plants in the stonecrop family, Crassulaceae, consisting of about ~70 taxa found in southwestern North America and Guadalupe Island. The genus is very morphologically diverse, with species ranging from large evergreen plants to tiny deciduous geophytic plants. The flowers of Dudleya have parts numbered in fives, with the petals arranged in tubular, star-shaped, or bell-shaped forms and, when fruiting, are filled with tiny, crescent-shaped seeds.

Description
Characteristics and subgenera This genus is the only taxon of the Sedeae tribe to have evolved sympodial branching. The genus is traditionally divided into three subgenera, two of which were formerly their own genera. The subgenera consist of Dudleya, Stylophyllum, and Hasseanthus. This waxy coating is usually white, chalky, and mealy, and is the namesake in many epithets, like chalk dudleya, powdery liveforever, and giant chalk dudleya. The wax on the giant chalk dudleya is a surface with some of the highest ultraviolet reflectivity ever discovered in plants. When exposed to water, the wax may coat drops on leaves, preventing their evaporation. The farina is mostly composed of pentacyclic triterpenoids. Other members, such as D. viscida and D. anomala, have a clear, sticky layer on their foliage. This adhesive layer ultimately helps the plant protect its leaves from the sun, much like ones with "white" wax. It accomplishes this solar protection by allowing dust to adhere to the sticky layer on the leaves, which coats the foliage with dust and debris, blocking solar radiation. == Taxonomy ==
Taxonomy
Taxonomic history Early history The earliest species of the genus described was Cotyledon caespitosa, by Adrian Hardy Haworth in 1803, which would later be known as Dudleya caespitosa. The same species would later be described again as Sedum cotyledon by Joseph Franz von Jacquin in 1811, and in 1840 Thomas Nuttall described both Echeveria pulverulenta (Dudleya pulverulenta) and Echeveria lanceolata (Dudleya lanceolata). The differing generic placements of these early descriptions suggest that taxonomic disagreements over the genus had an early start, with some of these generic placements persisting even after the proper description of the genus by Britton and Rose. In the 1930s, Alwin Berger revised the status of Dudleya and Stylophyllum into a sectional ranking within Echeveria, while merging the Hasseanthus genera into Sedum. This was in part due to a belief that Dudleya had evolved from Echeveria radiating northward from Mexico, evidenced by the similar tubular corollas, while Hasseanthus possessed aesthetic similarities to Sedum, primarily flowers with broadly spreading petals. Stylophyllum was believed to occupy a transitory position between Dudleya and Hasseanthus. These conclusions were widely accepted by botanists for the first half of the 20th century, until proper molecular and phylogenetic analyses began to appear. Modern classifications and phylogenetic research Around 1993, one of the only literary treatments of the genus was written, Paul H. Thomson's Dudleya and Hasseanthus Handbook. Research in the 21st century has supported the hypothesis of Dudleya's relation to Sedum. Phylogenetic analysis has given weight to this hypothesis, concluding that Dudleya is much closer to North American members of Sedoideae than to Echeverioideae. The plant Sedum spathulifolium, which co-exists with Dudleya in Oregon and California, has been shown in multiple phylogenetic reconstructions to be a sister taxon to Dudleya, although it may not be the true sister taxon, as larger samplings are needed. Berger's hypothesis of a relationship between Hasseanthus and Sedum may still prove true, as in addition to morphological similarities, the Hasseanthus-type taxa could be basal to the genus. However, no truly conclusive evidence of significant divergence within Dudleya exists, and it remains unclear which groups are basal or derived. Selected speciesDudleya abramsiiDudleya abramsii subsp. abramsii – Abrams' liveforever • Dudleya abramsii subsp. affinis – San Bernardino Mountains liveforever • Dudleya abramsii subsp. bettinae – Betty's dudleya • Dudleya abramsii subsp. calcicola – limestone dudleya • Dudleya abramsii subsp. murina – Mouse-gray dudleya • Dudleya abramsii subsp. setchellii – Santa Clara Valley dudleya • Dudleya acuminata – Vizcaino liveforever • Dudleya albiflora– White-flower liveforever • Dudleya anomala – Todos Santos liveforever • Dudleya anthonyi – Anthony's liveforever • Dudleya attenuata – Orcutt's liveforever, tapertip liveforever • Dudleya arizonica – Arizona chalk dudleya. Formerly a subspecies of Dudleya pulverulenta. • Dudleya blochmaniae – Blochman's liveforever, Blochman's dudleya • Dudleya brevifolia – Short-leaved liveforever, short-leaved dudleya. Formerly a subspecies of D. blochmaniae. • Dudleya brittonii – Britton's dudleya, giant chalk dudleya • Dudleya caespitosa – Coast dudleya, sea lettuce, sand lettuce • Dudleya campanulata – Punta Banda liveforever • Dudleya candelabrum – Candleholder liveforever • Dudleya candida – Coronados liveforever • Dudleya cedrosensisDudleya crassifolia – Thick-leaf dudleya • Dudleya cultrata – Knife-leaved liveforever, maritime succulent liveforever • Dudleya cymosa – Canyon liveforever • Dudleya cymosa subsp. agourensis – Agoura Hills dudleya • Dudleya cymosa subsp. crebrifolia – San Gabriel River dudleya • Dudleya cymosa subsp. costatifolia – Pierpoint Springs dudleya • Dudleya cymosa subsp. cymosa – Coast Range dudleya • Dudleya cymosa subsp. marcescens – Marcescent dudleya • Dudleya cymosa. subsp. ovatifolia – Santa Monica Mountains dudleya • Dudleya cymosa subsp. paniculata – Diablo range dudleya • Dudleya cymosa subsp. pumila – Transverse ranges liveforever • Dudleya densiflora – San Gabriel Mountains liveforever • Dudleya edulis – Fingertips, lady fingers, San Diego dudleya • Dudleya farinosa – Bluff lettuce, powdery liveforever, powdery dudleya • Dudleya formosa – La Mision liveforever • Dudleya gatesii – Gates' liveforever • Dudleya gnoma – Munchkin dudleya • Dudleya greenei – Greene's dudleya • Dudleya guadalupensis – Guadalupe liveforever • Dudleya ingens – Baja liveforever, rock liveforever • Dudleya hendrixii – Hendrix's liveforever • Dudleya lanceolata – Lanceleaf liveforever. • Dudleya linearis – San Benito Island liveforever • Dudleya multicaulis – Many-stemmed dudleya • Dudleya nubigena – Cape liveforever • Dudleya nesiotica – Santa Cruz Island liveforever • Dudleya pachyphytum – Cedros Island dudleya • Dudleya palmeri – Palmer's liveforever • Dudleya parva – Conejo dudleya • Dudleya pauciflora– Few-flower liveforever • Dudleya pulverulenta – Chalk lettuce, chalk dudleya, chalk liveforever • Dudleya rigida – Sierra de la Laguna liveforever • Dudleya rigidiflora – Playa Maria liveforever • Dudleya rubens – Red flowering liveforever • Dudleya saxosa – Panamint liveforever • Dudleya saxosa subsp. collomiae – Gila County liveforever • Dudleya saxosa subsp. aloides – Desert liveforever • Dudleya saxosa subsp. saxosa – Panamint liveforever • Dudleya stolonifera – Laguna Beach liveforever • Dudleya traskiae – Santa Barbara Island liveforever • Dudleya variegata – Variegated liveforever • Dudleya verityi – Verity's liveforever • Dudleya virens – Alabaster plant, Island liveforever • Dudleya viridicata – Colonet liveforever • Dudleya viridisDudleya viscida – Sticky dudleya Etymology The genus is named after William Russel Dudley, the first head of the botany department at Stanford University. The name of the subgenera Stylophyllum is an allusion to the pencil-shaped leaves of the type species, Stylophyllum edule, now known as Dudleya edulis.'' • Tiipai: milhka’mey (wide-leaf); milh kajmila (narrow-leaf) • Paipai: awi mielh == Reproductive biology ==
Reproductive biology
Pollinators and flower morphology The pollinators of Dudleya are mostly hummingbirds and bees, although this has been inferred in some species only from flower morphology. The genus exhibits a wide array of diverse flower varieties, even within species complexes. The flowers are mostly hermaphroditic, although some individuals may have sterile anthers. Plants with long, tubular, reddish flowers, mostly in the subgenus Dudleya, are adapted towards pollination via hummingbirds, while short, spreading, yellow flowers favor pollination by insects. Despite the evolution of long flowers towards hummingbirds, the long flowers also show no performance disadvantage in pollination environments exclusively composed of insects. While the long flowers may provide an evolutionary advantage for more effective pollination, more energy is required to produce the larger amounts of floral tissue. The flower morphology of Dudleya also corresponds with the aforementioned groups of Hasseanthus, Stylophyllum, and Dudleya. The flowers in Hasseanthus and Stylophyllum are broad, white to yellow, and spreading, with bumblebees and bees as primary pollinators, and hummingbirds as occasional pollinators. As these species grade towards the subgenus Dudleya, the petals begin to fuse at the base, with the flowers becoming tubular and red, due to an association with hummingbird pollinators. This shift in pollinators corresponds to larger nectar volumes and higher energy contents in accordance with the demands of hummingbird pollinators. File:2017-06-17-14.05.23 ZS PMax Dudleya caespitosa-1 - Flickr - John Rusk.jpg|subg. '' — The flowers of Dudleya caespitosa'' File:2017-08-19-13.55.44 ZS PMax Dudleya anthonyi—1—DxO (36281178490).jpg|subg. '' — The flowers of Dudleya anthonyi'' File:Dudleyaedulis.jpg|subg. '' — The flowers of Dudleya edulis'' File:Dudleya virens hassei.jpg|subg. '' — The flowers of Dudleya virens subsp. hassei'' File:Dudleyablochmaniae.jpg|subg. '' — The flowers of Dudleya blochmaniae'' File:Dudleya variegata 186339494.jpg|subg. '' - The flowers of Dudleya variegata'' == Distribution and habitat ==
Distribution and habitat
Dudleya species are widespread and diverse in their range, but are typically found in rock outcroppings, cliff faces, or road cuts, where their leaves help them store water in a setting too dry for most types of plants. Most are small and inconspicuous when not in bloom. The diversity of species of Dudleya is centered in Southern California and northern Baja California. '', found on the Baja California peninsula ==Horticulture==
Horticulture
Watering In horticulture, Dudleya should be planted at an angle. This allows accumulated water to drain from the nestlike center of the plant, thus preventing microbial decay. Dudleya should not be watered from directly above, as this may damage their chalky coating known as farina, which is present on numerous species. Clay is preferable to plastic pots when growing the plants due to the advantages with water drainage. Plastic pots may also kill the plants by overheating the roots. Powdery mildew has also been recorded in cultivated Dudleya. Shade The amount of shade a species of Dudleya requires is dependent on the location. Plants growing in coastal regions may require little shade at all, whilst plants growing in the deserts, inland valleys and mountains will require shade. The majority of plants in the genus will appreciate north-facing sites and shade during the heat of the day. During the summer months, 50% shade may be beneficial for plants. If a cold-tolerant Dudleya is grown during a freeze or snow, it should be shaded as not to damage the plant, as a quick thaw may be detrimental. Asexual reproduction Vegetative reproduction may occur in multiple ways: Most Dudleya will not reproduce from leaves, with the leaves simply dying after being severed. • Plant tissue culture methods are also available, and becoming more advanced. Currently, plant tissue culture is used for Dudleya propagation in commercial and conservation settings. == Ethnobotany ==
Ethnobotany
Kumeyaay and Paipai The indigenous peoples of the Kumeyaay and Paipai region utilized the genus for both medicinal and agricultural purposes. The tender, succulent leaves were chewed on to alleviate thirst, or used to treat calluses and corns. The budding inflorescences, in their early stages, were used as food, with a sweet flavor and juicy texture. The roots were pounded up and soaked in water, used as an astringent to "tighten the gums." The roots were also boiled whole as a decoction for asthma. == Conservation ==
Conservation
Several species of Dudleya are threatened by urban development in Coastal California and Mexico, and anthropogenic-induced wildfires. However, one of the most critical threats to Dudleya species is poaching, partially caused by a demand from East Asian succulent collectors paying lucrative prices for certain Dudleya species.'', which was threatened by poachers The population of Verity's liveforever (Dudleya verityi), which was nearly wiped out during the 2013 Springs Fire, was targeted by poachers. The Cedros Island liveforever (Dudleya pachyphytum) is a rare and extremely specialized Dudleya endemic to Cedros Island. In 2016, Korean nationals began moving to Bahia Tortugas, a locality in Baja California Sur, to facilitate the poaching and shipping of the plants. The species was seriously threatened after Mexican soldiers discovered poachers taking nearly 5,000 rosettes in a tractor-trailer. It was suspected the poachers rappelled onto the succulent's location via helicopter, as D. pachyphytum occupies a nearly inaccessible habitat. In 2019, the deaths and injuries of several fishermen from Bahía Tortugas who were on Cedros Island was allegedly the result of Dudleya trafficking, a conflict with the Sinaloa Cartel, or both. In 2020, the Mexican Navy in the Second Naval Region revealed that two fishermen were killed after a dispute emerged over the trafficking of the rare plant. Bluff lettuce (Dudleya farinosa) was also targeted by poachers in numerous large-scale operations. Although not particularly rare, the size of the poaching operation pose a serious ecological threat. Starting in 2017, the U.S. Customs and Border Protection, along with the California Department of Fish and Wildlife, discovered large amounts of D. farinosa being shipped out of the country. South Korean and Chinese nationals have both been arrested in the smuggling of D. farinosa. According to nursery owners responsible for legally exporting Dudleya, buyers in Asia desired plants directly from the wild, owing to the aesthetic quality of their long caudices and weathered leaves. The candleholder dudleya (Dudleya candelabrum), native to the northern Channel Islands, was reported to have been poached, with the thieves shipping plants to South Korea. In response to the poaching of Dudleya, California State Assembly member Chris Ward proposed Bill AB-223, sponsored by the California Native Plant Society, which would make it illegal to poach Dudleya from state or private lands without a permit. The California Native Plant Society and conservationists have also initiated propagation programs to oversaturate the market as a means to deter poachers. On September 28, 2021, governor Gavin Newsom signed AB-223 into law. == See also ==
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