Dudleya Greenei
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''Dudleya'', commonly known as liveforevers, is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
rosette Rosette is the French diminutive of ''rose''. It may refer to: Flower shaped designs * Rosette (award), a mark awarded by an organisation * Rosette (design), a small flower design *hence, various flower-shaped or rotational symmetric forms: ** R ...
-forming
succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
plants in the stonecrop family,
Crassulaceae The Crassulaceae (, from Latin ''crassus'', thick), also known as the crassulas, the stonecrops or the orpine family, are a diverse Family (biology), family of dicotyledon angiosperms primarily characterized by succulent leaves and a form of phot ...
, consisting of about 68 taxa in southwestern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
and
Guadalupe Island Guadalupe Island () is a volcanic island located off the western coast of Mexico's Baja California peninsula and about southwest of the city of Ensenada in the state of Baja California, in the Pacific Ocean. The various volcanoes are extinc ...
. The species come in many forms, some large and
evergreen In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
, others
geophytic A storage organ is a part of a plant specifically modified for storage of energy (generally in the form of carbohydrates) or water. Storage organs often grow underground, where they are better protected from attack by herbivores. Plants that have ...
and
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
. 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. The genus evolved as neoendemics, from ancestors in the stonecrop genus, ''
Sedum ''Sedum'' is a large genus of flowering plants in the family Crassulaceae, members of which are commonly known as stonecrops. The genus has been described as containing up to 600 species, subsequently reduced to 400–500. They are leaf succule ...
''. The ancestors radiated southward from ''Sedum'' during the creation of the dry summer climate, in the California region, five million years ago. Early botanists classified the larger species as ''
Echeveria ''Echeveria'' is a large genus of flowering plants in the family (biology), family Crassulaceae, native plant, native to semi-desert areas of Central America, Mexico and northwestern South America. Description Echeveria plants are evergreen. Flo ...
'' and ''
Cotyledon A cotyledon ( ; ; "a cavity, small cup, any cup-shaped hollow", gen. (), ) is a "seed leaf" – a significant part of the embryo within the seed of a plant – and is formally defined as "the embryonic leaf in seed-bearing plants, one or mor ...
'', while the geophytic species were placed under ''Sedum''. Taxonomic efforts, started by
Joseph Nelson Rose Joseph Nelson Rose (January 11, 1862 – May 4, 1928) was an American botanist. He was born in Union County, Indiana. His father died serving during the Civil War when Joseph Rose was a young boy. He later graduated from high school in Liberty, ...
and
Nathaniel Lord Britton Nathaniel Lord Britton (1859 – 1934) was an American botanist and taxonomist who co-founded the New York Botanical Garden in the Bronx, New York (state), New York. Early life Britton was born on the 15 of January 1859 at New Dorp, Staten Island ...
, created three genera; these initial genera were all eventually subsumed into ''Dudleya'', proper, following
Reid Moran Reid Venable Moran (June 30, 1916 – January 21, 2010) was an American botanist and the curator of botany at the San Diego Natural History Museum from 1957 to 1982. Moran was the world authority on the Crassulaceae, a family of succulent plants, ...
's investigations into the genus.
Phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
research is still at an early stage in the genus, and is complicated by the fact that many species are becoming
endangered An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching, inv ...
and over-harvested (poached). ''Dudleya'' is a relatively obscure genus, in comparison to other, more widely-cultivated succulents; converging interests, by succulent collectors,
native plant In biogeography, a native species is indigenous to a given region or ecosystem if its presence in that region is the result of only local natural evolution (though often popularised as "with no human intervention") during history. The term is equi ...
enthusiasts and gardeners alike, have led to the wider cultivation of many species as
ornamental plant Ornamental plants or ''garden plants'' are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars th ...
s. In the wild, many species of ''Dudleya'' are vulnerable, as
land development Land development is the alteration of landscape in any number of ways, such as: * Changing landforms from a natural or semi-natural state for a purpose such as agriculture or House, housing * subdivision (land), Subdividing real estate into Lot ( ...
and
poachers Poaching is the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set against the hunti ...
often threaten particularly
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development and growth *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ec ...
populations of plants. Poached plants are often shipped to
East Asia East Asia is a geocultural region of Asia. It includes China, Japan, Mongolia, North Korea, South Korea, and Taiwan, plus two special administrative regions of China, Hong Kong and Macau. The economies of Economy of China, China, Economy of Ja ...
, especially
South Korea South Korea, officially the Republic of Korea (ROK), is a country in East Asia. It constitutes the southern half of the Korea, Korean Peninsula and borders North Korea along the Korean Demilitarized Zone, with the Yellow Sea to the west and t ...
, where hybridisation and cultivation of succulents is very popular.
Conservationists The conservation movement, also known as nature conservation, is a political, environmental, and social movement that seeks to manage and protect natural resources, including animal, fungus, and plant species as well as their habitat for the ...
, nurseries and
government A government is the system or group of people governing an organized community, generally a State (polity), state. In the case of its broad associative definition, government normally consists of legislature, executive (government), execu ...
s combat ''Dudleya'' poaching through propagation programs and protection laws.


Description


Characteristics and subgenera

This genus is the only taxon of the Sedeae tribe to have evolved
sympodial branching In botany, sympodial growth is a bifurcating branching pattern where one branch develops more strongly than the other, resulting in the stronger branches forming the primary shoot and the weaker branches appearing laterally. A sympodium, also ...
. The genus is traditionally divided into three subgenera, two of which were formerly their own genera. The subgenera consist of ''Dudleya, Stylophyllum'', and ''Hasseanthus''.


Epicuticular wax

One of the most famous features of some ''Dudleya'' is their waxy coating. Numerous species have their foliage covered in an
epicuticular wax Epicuticular wax is a waxy coating which covers the outer surface of the plant cuticle in land plants. It may form a whitish film or bloom on leaves, fruits and other plant organs. Chemically, it consists of hydrophobic organic compounds, mainly ...
, or farina. 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 Triterpenes are a class of terpenes composed of six isoprene units with the molecular formula C30H48; they may also be thought of as consisting of three terpene units. Animals, plants and fungi all produce triterpenes, including squalene, the pre ...
. Other members, such as '' D. viscida'' and '' D. anomala'', have a clear, sticky layer on their foliage.Dudleya anomala ''(Davidson) Moran''. Desert Plant Life. 14: 191. 1942. 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


Taxonomic history


Early history

The earliest species of the genus described was ''Cotyledon caespitosa'', by
Adrian Hardy Haworth Adrian Hardy Haworth (19 April 1767, in Kingston upon Hull, Hull – 24 August 1833, in Chelsea, London, Chelsea) was an England, English entomologist, botanist and carcinologist. Family The younger son of Benjamin Haworth, of Haworth Hall and ...
in 1803, which would later be known as ''
Dudleya caespitosa ''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, t ...
.'' The same species would later be described again as ''Sedum cotyledon'' by
Joseph Franz von Jacquin Joseph "Krystel" Franz Freiherr von Jacquin or Baron Joseph von Jacquin (7 February 1766, in Schemnitz (now Banská Štiavnica) – 26 October 1839, in Vienna) was an Austrian scientist who studied medicine, chemistry, zoology and botany. ...
in 1811, and in 1840
Thomas Nuttall Thomas Nuttall (5 January 1786 – 10 September 1859) was an English botanist and zoologist who lived and worked in America from 1808 until 1841. Nuttall was born in the village of Long Preston, near Settle in the West Riding of Yorkshire a ...
described both ''Echeveria pulverulenta'' (''
Dudleya pulverulenta ''Dudleya pulverulenta'' is a species of Perennial plant, 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 greatl ...
'') 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.
Nathaniel Lord Britton Nathaniel Lord Britton (1859 – 1934) was an American botanist and taxonomist who co-founded the New York Botanical Garden in the Bronx, New York (state), New York. Early life Britton was born on the 15 of January 1859 at New Dorp, Staten Island ...
and
Joseph Nelson Rose Joseph Nelson Rose (January 11, 1862 – May 4, 1928) was an American botanist. He was born in Union County, Indiana. His father died serving during the Civil War when Joseph Rose was a young boy. He later graduated from high school in Liberty, ...
's revision of the North American
Crassulaceae The Crassulaceae (, from Latin ''crassus'', thick), also known as the crassulas, the stonecrops or the orpine family, are a diverse Family (biology), family of dicotyledon angiosperms primarily characterized by succulent leaves and a form of phot ...
upended many of the early taxonomic classifications, with the newly-minted genus ''Dudleya'' containing 60 species, of which 41 were newly described by Britton and Rose. The two also defined the related genera ''Stylophyllum'', which contained 12 species, and ''Hasseanthus'', with 4 species. The primary differences between ''Dudleya'' and ''Stylophyllum'' were between the leaf shape, floral structure and petal orientation, whilst ''Hasseanthus'' was characterized by a different vegetative structure and more niche adaptations, primarily corm-like stems and deciduous leaves.Uhl, C. H., & Moran, R. (1953). The cytotaxonomy of Dudleya and Hasseanthus. ''American Journal of Botany'', 492-502. In the 1930s,
Alwin Berger Alwin Berger (28 August 1871 – 20 April 1931) was a German botanist best known for his contribution to the nomenclature of succulent plants, particularly agaves and cacti. Born in Germany he worked at the botanical gardens in Dresden and Fra ...
revised the status of ''Dudleya'' and ''Stylophyllum'' into a sectional ranking within ''
Echeveria ''Echeveria'' is a large genus of flowering plants in the family (biology), family Crassulaceae, native plant, native to semi-desert areas of Central America, Mexico and northwestern South America. Description Echeveria plants are evergreen. Flo ...
'', while merging the ''Hasseanthus'' genera into ''
Sedum ''Sedum'' is a large genus of flowering plants in the family Crassulaceae, members of which are commonly known as stonecrops. The genus has been described as containing up to 600 species, subsequently reduced to 400–500. They are leaf succule ...
''. 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.


Reid Moran, Charles H. Uhl and early phylogenetics

In 1942,
Reid Moran Reid Venable Moran (June 30, 1916 – January 21, 2010) was an American botanist and the curator of botany at the San Diego Natural History Museum from 1957 to 1982. Moran was the world authority on the Crassulaceae, a family of succulent plants, ...
, a botanist with a longstanding interest in the genera, offered a new revision of the taxa. Moran recognized ''Dudleya'' as being distinct from ''Echeveria'', and merged ''Stylophyllum'' as a subgenera along with ''Eududleya'' (which was changed to just subgenus ''Dudleya''). Moran still recognized ''Hasseanthus'' as a separate genus, but realized that it was closely related to ''Dudleya''. With assistance from cytologist Charles H. Uhl, Moran came to the conclusion that ''Dudleya'' and ''Hasseanthus'' were related closer to each other than they were to ''Echeveria'' or ''Sedum''. As Moran and Uhl conducted more cytological and taxonomic research on the genus, it became clear that ''Hasseanthus'' was also a subgenera of ''Dudleya'', citing the karyological uniformity and the formation of hybrids between the genera. In contrast, while ''Echeveria'' and ''Sedum'' freely hybridize within themselves and each other, all attempts at hybridizing ''Dudleya'' with ''Echeveria'' have failed. Uhl came to the conclusion that there are probably no intergeneric hybrids with ''Dudleya'', believing that ''Dudleya'' had split from ''Echeveria'' far enough that intergeneric reproductive success was unlikely. Despite Uhl's research, he still believed that ''Dudleya'' had evolved northward from Mexico with ''Echeveria'', probably some time before the formation of the summer-dry climate of California, making ''Dudleya'' a
paleoendemic Paleoendemism along with neoendemism is a possible subcategory of endemism. Paleoendemism refers to species that were formerly widespread but are now restricted to a smaller area. Neoendemism refers to species that have recently arisen, such as th ...
.


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''. In the book, Thompson made numerous changes and adjustments to species, and described several new species. The book included a large number of photographs, extensive descriptions, and horticultural practices for the care of ''Dudleya,'' derived from a long interest growing, visiting and collecting the plants. However, he failed to follow the
International Code of Botanical Nomenclature The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN or ICNafp) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all tho ...
guidelines for describing new taxa, forgoing designated
type specimens In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes t ...
, which invalidated his treatment. His treatment also contains extensive
pseudoscientific Pseudoscience consists of statements, beliefs, or practices that claim to be both scientific and factual but are incompatible with the scientific method. Pseudoscience is often characterized by contradictory, exaggerated or unfalsifiable cl ...
material, including references to the lost continents of
Atlantis Atlantis () is a fictional island mentioned in Plato's works '' Timaeus'' and ''Critias'' as part of an allegory on the hubris of nations. In the story, Atlantis is described as a naval empire that ruled all Western parts of the known world ...
,
Lemuria Lemuria (), or Limuria, was a continent proposed in 1864 by zoologist Philip Sclater, theorized to have sunk beneath the Indian Ocean, later appropriated by occultists in supposed accounts of human origins. The theory was discredited with the dis ...
and Mu, a belief that
cosmic ray Cosmic rays or astroparticles are high-energy particles or clusters of particles (primarily represented by protons or atomic nuclei) that move through space at nearly the speed of light. They originate from the Sun, from outside of the ...
s and
divine intervention Divine intervention is an event that occurs when a deity (i.e. God or gods) becomes actively involved in changing some situation in human affairs. In contrast to other kinds of divine action, the expression "divine ''intervention''" implies that ...
directly induced the chromosome counts and
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
of ''Hasseanthus'' and ''Dudleya'', and criticism of
paleontologists Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
and the scientifically accepted
age of the Earth The age of Earth is estimated to be 4.54 ± 0.05 billion years. This age may represent the age of Earth's accretion (astrophysics), accretion, or Internal structure of Earth, core formation, or of the material from which Earth formed. This dating ...
. Later research challenged the notion that ''Dudleya'' and ''Echeveria'' formed two distinct lineages within Echeverioideae. Joachim Thiede instead proposed that ''Dudleya'' evolved completely independently from ''Echeveria'', evolving from relatives within the ''Leucosedum'' clade like '' Sedum moranii''. This places ''Dudleya'' as a
neoendemic Neoendemism is one of two sub-categories of endemism, the ecological state of a species being unique to a defined geographic location. Specifically, neoendemic species are those that have recently arisen, through divergence and reproductive isolati ...
that evolved recently with the formation of the summer-dry climate in California. This was evidenced by the similarities of seed-surface ornamentation between ''Dudleya'' and ''Sedum'', which is unlike ''Echeveria'', and the reproductive isolation between ''Echeveria'' and ''Dudleya''.Thiede, J., & Eggli, U. (2007). Crassulaceae. In ''Flowering Plants· Eudicots'' (pp. 83-118). Springer, Berlin, Heidelberg. Chicago 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 ''Sedum spathulifolium'' is a species of flowering plant in the family Crassulaceae known by the common names broadleaf stonecrop, yellow stonecrop, and spoon-leaved stonecrop. An evergreen perennial, it is native to western North America from Br ...
'', 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 species

*''
Dudleya abramsii ''Dudleya abramsii'' is a species complex of succulent plants native to California and parts of Baja California. There are numerous subspecies, some critically endangered, with varying habits and lifestyles, but most often characterized by a smal ...
'' ** ''Dudleya 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 ''Dudleya blochmaniae'' is a summer-deciduous succulent plant known by the common names Blochman's liveforever or Blochman's dudleya. This species of ''Dudleya'' survives part of the year with no aboveground presence, surviving as underground cor ...
'' – Blochman's liveforever, Blochman's dudleya *''
Dudleya brevifolia ''Dudleya brevifolia'', is a rare succulent plant known by the common name short-leaved liveforever, short-leaved dudleya or rarely the Del Mar Hasseanthus. It is an edaphic Endemism, endemic that only grows on Lindavista formation Raised beach, ...
'' – Short-leaved liveforever, short-leaved dudleya. Formerly a subspecies of ''D. blochmaniae''. *''
Dudleya brittonii ''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, so ...
'' – Britton's dudleya, giant chalk dudleya *''
Dudleya caespitosa ''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, t ...
'' – Coast dudleya, sea lettuce, sand lettuce *'' Dudleya campanulata'' – Punta Banda liveforever *'' Dudleya candelabrum'' – Candleholder liveforever *'' Dudleya candida'' – Coronados liveforever *'' Dudleya cedrosensis'' *'' Dudleya crassifolia'' – Thick-leaf dudleyaDodero, M. W. and M. G. Simpson. (2012)
"''Dudleya crassifolia'' (Crassulaceae), a new species from northern Baja California, Mexico"
''Madroño'' 59(4) 223–229.
*'' 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 ''Dudleya formosa'', known by the common name La Misión liveforever, is a species of Perennial plant, perennial Succulent plant, succulent plant Endemism, endemic to the Valle de Guadalupe, Guadalupe Valley in Baja California. It is characterize ...
'' – 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 ''Dudleya pulverulenta'' is a species of Perennial plant, 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 greatl ...
'' – 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 ''Dudleya verityi'' is a rare species of succulent plant known by the common name Verity's liveforever. It is Endemism, endemic to Ventura County, California, where it is known from only three occurrences in the vicinity of Conejo Mountain betwee ...
'' – Verity's liveforever *'' Dudleya virens'' – Alabaster plant, Island liveforever *'' Dudleya viridicata'' – Colonet liveforever *'' Dudleya viridis'' *'' Dudleya viscida'' – Sticky dudleya


Etymology

The genus is named after William Russel Dudley, the first head of the botany department at
Stanford University Leland Stanford Junior University, commonly referred to as Stanford University, is a Private university, private research university in Stanford, California, United States. It was founded in 1885 by railroad magnate Leland Stanford (the eighth ...
. The term ''liveforever'' may refer to the longevity and hardiness of some plants in the wild. The name of the subgenera ''Stylophyllum'' is an
allusion Allusion, or alluding, is a figure of speech that makes a reference to someone or something by name (a person, object, location, etc.) without explaining how it relates to the given context, so that the audience must realize the connection in the ...
to the pencil-shaped leaves of the type species, ''Stylophyllum edule'', now known as ''Dudleya edulis''. The name of the subgenera ''Hasseanthus'' was in honor of Dr. Hermann E. Hasse, a surgeon and collector of California flora. ''Dudleya virens'' subsp. ''hassei'' is also named after him.


Nomenclature

*
English English usually refers to: * English language * English people English may also refer to: Culture, language and peoples * ''English'', an adjective for something of, from, or related to England * ''English'', an Amish ter ...
: ''Dudleya, liveforever'' *
Spanish Spanish might refer to: * Items from or related to Spain: **Spaniards are a nation and ethnic group indigenous to Spain **Spanish language, spoken in Spain and many countries in the Americas **Spanish cuisine **Spanish history **Spanish culture ...
: s''iempreviva, mezcalito'' * Tiipai: ''milhka’mey'' (wide-leaf); ''milh kajmila'' (narrow-leaf) * Paipai: ''awi mielh''


Reproductive biology


Pollinators and flower morphology

The pollinators of ''Dudleya'' are mostly
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
s and
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
s, 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 complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
es. The flowers are mostly
hermaphroditic A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic. The individuals of many ...
, 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.


Seeds

Seed morphology also differs between ''Dudleya''. Larger ''Dudleya'' have noticeably smaller seeds; this is likely a result of an evolutionary tradeoff. The investment in growing large allows plants to retain moisture in periods of drought, at the cost of energy placed into reproduction. Other reasons for smaller seeds may also be due to allocation of resources into vegetative reproduction via pup rosettes, instead of sexual reproduction. In contrast, the small, inconspicuous, ''Hasseanthus''-type members produce the largest seeds in the genus. These larger seeds are not distributed very far, which contributes to the restricted ranges of the ''Hasseanthus'' subgenus.Amoroso, D. M., & Wilson, P. (2018). Ten Cases of Divergence in the Seedling Ecology of Dudleya (Crassulaceae). ''Systematic Botany'', ''43''(4), 889-900. 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

''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 two predominant habitats where plants of this genera may be found are by the coast or mountains. Dudleya favor moderate temperatures, summer dormancy, winter precipitation, and rocky habitats, which means they may be found in diverse, disjunct locales from oceanic bluffs on the California coast to sky islands in Arizona. Regions where ''Dudleya'' can be found include
The Californias The Californias (), occasionally known as the Three Californias or the Two Californias, are a region of North America spanning the United States and Mexico, consisting of the U.S. state of California and the Mexican states of Baja California a ...
,
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
, coastal
Sonora Sonora (), officially Estado Libre y Soberano de Sonora (), is one of the 31 states which, along with Mexico City, comprise the Administrative divisions of Mexico, Federal Entities of Mexico. The state is divided into Municipalities of Sonora, 72 ...
and
Oregon Oregon ( , ) is a U.S. state, state in the Pacific Northwest region of the United States. It is a part of the Western U.S., with the Columbia River delineating much of Oregon's northern boundary with Washington (state), Washington, while t ...
, and southern
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
and
Nevada Nevada ( ; ) is a landlocked state in the Western United States. It borders Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the seventh-most extensive, th ...
.Thiede, J. (2004). The genus Dudleya Britton & Rose (Crassulaceae): its systematics and biology. ''Cactus and Succulent Journal (US)'', ''76'', 224-231. The diversity of species of ''Dudleya'' is centered in
Southern California Southern California (commonly shortened to SoCal) is a geographic and Cultural area, cultural List of regions of California, region that generally comprises the southern portion of the U.S. state of California. Its densely populated coastal reg ...
and northern
Baja California Baja California, officially the Free and Sovereign State of Baja California, is a state in Mexico. It is the northwesternmost of the 32 federal entities of Mexico. Before becoming a state in 1952, the area was known as the North Territory of B ...
.


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. During the late fall to early spring period of growing, watering should be done frequently in order to promote good growth and flower development. Plants grown with exposure to rainfall, provided they are in a Mediterranean climate, they may obtain optimum moisture. In contrast, plants grown indoors or in greenhouses should be watered to ensure that the plant maximizes growth. During the dormant period, from late spring to fall, watering should be limited to bi-weekly intervals, but not completely cut out. In nature, plants may completely desiccate themselves during the dormant period. Deciduous members of the genus in the ''Hasseanthus'' grouping should not be watered at all during dormancy.


Growing medium

Growing mediums should attempt to imitate the preferred soil of each species in nature. The most essential element for the medium is good drainage. Good drainage is important, as ''Dudleya'' are very susceptible to mold and fungi. How quickly the medium drains should be determined by the amount of shade in the location, which affects the rate of evaporation that the soil will go through. Thus, well-shaded ''Dudleya'' must have excellent drainage. Fertilizers may be used to maintain good color and growth, but they should be diluted. 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.
Mealybug Mealybugs are insects in the family Pseudococcidae, unarmored scale insects found in moist, warm habitats. Of the more than 2,000 described species, many are considered pests as they feed on plant juices of greenhouse plants, house plants and ...
s and
aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...
s are main pests of ''Dudleya''.
Powdery mildew Powdery mildew is a fungus, fungal disease that affects a wide range of plants. Powdery mildew diseases are caused by many different species of Ascomycota, ascomycete fungi in the order Erysiphales. Powdery mildew is one of the easier plant disea ...
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.


Propagation

Unlike their related genera ''Echeveria'', many ''Dudleya'' cannot be propagated through leaf cuttings. Propagation is mostly achieved via offsets,
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ...
by seed, or in nurseries, plant tissue culture.


Sexual reproduction

''Dudleya'' seeds are crescent shaped and near-microscopic, and can be collected by taking the dried flower buds and crushing the ovules inside, which can contain hundreds of seeds. Seeds can be sown into a soilless medium, such as
pumice Pumice (), called pumicite in its powdered or dust form, is a volcanic rock that consists of extremely vesicular rough-textured volcanic glass, which may or may not contain crystals. It is typically light-colored. Scoria is another vesicula ...
or
vermiculite Vermiculite is a hydrous phyllosilicate mineral which undergoes significant expansion when heated. Exfoliation occurs when the mineral is heated sufficiently; commercial furnaces can routinely produce this effect. Vermiculite forms by the weathe ...
, after which germination can occur within a week to 10 days.


Asexual reproduction

Vegetative reproduction Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is a form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or specializ ...
may occur in multiple ways: # The rosette of the plant divides into two, eventually branching off to develop two separate rosettes. # Plantlets will form in the
axillary bud The axillary bud (or lateral bud) is an Plant embryogenesis, embryonic or Organogenesis#Plant organogenesis, organogenic Shoot (botany), shoot located in the axil of a leaf. Each bud has the potential to form shoots, and may be specialized in prod ...
s of old leaves around the primary rosettes, forming as many as 10 new rosettes. # After a
wildfire A wildfire, forest fire, or a bushfire is an unplanned and uncontrolled fire in an area of Combustibility and flammability, combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identified as a ...
, when the plant has its main rosettes burned off, new ones will emerge from either the stem or the roots, forming a new plant within a single growing season. # Rosettes emerge out of a long, horizontal root. This is commonplace when the plant has been decapitated on a cliff face, leaving only the long roots, which send out rosettes. # Plants multiply via stoloniferous growth. Many ''Dudleya'' may send out stoloniferous rosettes, but they do not root. Only does the Laguna Beach Liveforever (''Dudleya stolonifera'') sprout roots out of stoloniferous plantlets, yet this can be difficult or impossible to induce in captivity. # When the underground portion of the plant is lost to root rot, but the rosette and part of the caudex survive, the parts above the decayed roots can be removed cleanly, and planted provided they send out roots. Conversely, should the top of the rosette die but the lower stem still live, the deceased portion may be removed, a depression will remain in the caudex where the decayed rosette was removed, and at the edges new buds will sprout. # Propagation via leaves. Only certain species, like those of the ''Hasseanthus'' subgenus and others like '' Dudleya parva'', will root. Most ''Dudleya'' will not reproduce from leaves, with the leaves simply dying after being severed. #
Plant tissue culture Plant tissue culture is a collection of techniques used to maintain or grow plant cells, tissues, or organs under sterile conditions on a nutrient culture medium of known composition. It is widely used to produce clones of a plant in a method know ...
methods are also available, and becoming more advanced. Currently, plant tissue culture is used for ''Dudleya'' propagation in commercial and conservation settings.


Ethnobotany


Kumeyaay and Paipai

The indigenous peoples of the
Kumeyaay The Kumeyaay, also known as 'Iipai-Tiipai or by the historical Spanish name Diegueño, is a tribe of Indigenous peoples of the Americas who live at the northern border of Baja California in Mexico and the southern border of California in the Uni ...
and Paipai region utilized the genus for both
medicinal Medicine is the science and Praxis (process), practice of caring for patients, managing the Medical diagnosis, diagnosis, prognosis, Preventive medicine, prevention, therapy, treatment, Palliative care, palliation of their injury or disease, ...
and
agricultural Agriculture encompasses crop and livestock production, aquaculture, and forestry for food and non-food products. Agriculture was a key factor in the rise of sedentary human civilization, whereby farming of domesticated species created f ...
purposes. The tender,
succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
leaves were chewed on to alleviate
thirst Thirst is the craving for potable fluids, resulting in the basic instinct of animals to drink. It is an essential mechanism involved in fluid balance. It arises from a lack of fluids or an increase in the concentration of certain osmolites, suc ...
, or used to treat
callus A callus (: calluses) is an area of thickened and sometimes hardened skin that forms as a response to repeated friction, pressure, or other irritation. Since repeated contact is required, calluses are most often found on the feet and hands, b ...
es and
corns Corn most often refers to maize, the yellow, large-grained crop native to the Americas. It can also refer to the main cereal crop of a country or region: * Wheat, barley and oats in England and Wales * Oats in Scotland and Ireland Corn may also ...
. The budding
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
s, 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 An astringent (sometimes called adstringent) is a chemical that shrinks or constricts body tissues. The word derives from the Latin '' adstringere'', which means "to bind fast". Astringency, the dry, puckering or numbing mouthfeel caused by t ...
to "tighten the gums." The roots were also boiled whole as a decoction for asthma.Wilken, Michael A. (2012) ''An Ethnobotany of Baja California's Kumeyaay Indians.'' Retrieved 13 October 2021


Conservation

Several species of ''Dudleya'' are threatened by urban development in
Coastal California Coastal California, also known as the California Coastline and the Golden Coast, refers to the coastal regions of the U.S. state of California. The term is not primarily geographical as it also describes an area distinguished by cultural, economic ...
and Mexico, and anthropogenic-induced wildfires. However, one of the most critical threats to ''Dudleya'' species is
poaching Poaching is the illegal hunting or capturing of wild animals, usually associated with land use rights. Poaching was once performed by impoverished peasants for subsistence purposes and to supplement meager diets. It was set against the huntin ...
, partially caused by a demand from
East Asia East Asia is a geocultural region of Asia. It includes China, Japan, Mongolia, North Korea, South Korea, and Taiwan, plus two special administrative regions of China, Hong Kong and Macau. The economies of Economy of China, China, Economy of Ja ...
n
succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
collectors Collector(s) may refer to: Arts and entertainment * Collector (2011 film), ''Collector'' (2011 film), an Indian Malayalam film * Collector (2016 film), ''Collector'' (2016 film), a Russian film * Collectors (film), ''Collectors'' (film), a 2020 So ...
paying lucrative prices for certain ''Dudleya'' species. The population of Verity's liveforever (''Dudleya verityi''), which was nearly wiped out during the 2013
Springs Fire The Springs Fire was a wildfire in Ventura County, California in May 2013. Although the fire burned only 15 homes, it threatened 4,000. This threat passed when rain shower from few narrow cold-frontal rainbands moved through the California area ...
, was targeted by poachers. The Cedros Island liveforever (''Dudleya pachyphytum'') is a rare and extremely specialized ''Dudleya'' endemic to
Cedros Island Cedros Island (''Isla de Cedros'', "island of cedars" in Spanish (language), Spanish) is an island in the Pacific Ocean belonging to the state of Baja California, Mexico. The dry and rocky island had a population of 1,350 in 2005 and has an area o ...
. In 2016, Korean nationals began moving to Bahia Tortugas, a locality in
Baja California Sur Baja California Sur, officially the Free and Sovereign State of Baja California Sur, is a state in Mexico. It is the 31st and last state to be admitted, in 1974. It is also the second least populated Mexican state and the ninth-largest state by ...
, 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 The Sinaloa Cartel (, , after the native Sinaloa region), also known as the ''CDS'', the ''Guzmán-Loera Organization'', the ''Federation'', the ''Sinaloa Cartel'', or the Pacific Cartel, is a large, drug trafficking transnational organized cri ...
, or both. In 2020, the
Mexican Navy The Mexican Navy () is one of the components of the Mexican Armed Forces. The Secretariat of the Navy is in charge of administration of the navy. The commander of the navy is the Secretary of the Navy, who is both a cabinet minister and a career ...
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 United States Customs and Border Protection (CBP) is the largest federal law enforcement agency of the United States Department of Homeland Security. It is the country's primary border control organization, charged with regulating and facilita ...
, along with the
California Department of Fish and Wildlife The California Department of Fish and Wildlife (CDFW), formerly known as the California Department of Fish and Game (CDFG), is an American state agency under the California Natural Resources Agency. The Department of Fish and Wildlife manages ...
, discovered large amounts of ''D.'' ''farinosa'' being shipped out of the country.
South Korea South Korea, officially the Republic of Korea (ROK), is a country in East Asia. It constitutes the southern half of the Korea, Korean Peninsula and borders North Korea along the Korean Demilitarized Zone, with the Yellow Sea to the west and t ...
n and
Chinese Chinese may refer to: * Something related to China * Chinese people, people identified with China, through nationality, citizenship, and/or ethnicity **Han Chinese, East Asian ethnic group native to China. **'' Zhonghua minzu'', the supra-ethnic ...
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 Aesthetics (also spelled esthetics) is the branch of philosophy concerned with the nature of beauty and taste, which in a broad sense incorporates the philosophy of art.Slater, B. H.Aesthetics ''Internet Encyclopedia of Philosophy,'' , acces ...
quality of their long
caudices A caudex (: caudices) of a plant is a stem, but the term is also used to mean a rootstock and particularly a basal stem structure from which new growth arises.pages 456 and 695 In the strict sense of the term, meaning a stem, "caudex" is most o ...
and weathered leaves. The candleholder dudleya (''Dudleya candelabrum''), native to the northern
Channel Islands The Channel Islands are an archipelago in the English Channel, off the French coast of Normandy. They are divided into two Crown Dependencies: the Jersey, Bailiwick of Jersey, which is the largest of the islands; and the Bailiwick of Guernsey, ...
, was reported to have been poached, with the thieves shipping plants to South Korea. In response to the poaching of ''Dudleya,''
California State Assembly The California State Assembly is the lower house of the California State Legislature (the upper house being the California State Senate). The Assembly convenes, along with the State Senate, at the California State Capitol in Sacramento, Califor ...
member Chris Ward proposed Bil
AB-223
sponsored by the
California Native Plant Society The California Native Plant Society (CNPS) is a California environmental non-profit organization (501(c)(3)) that seeks to increase understanding of California's native flora and to preserve it for future generations. The mission of CNPS is to c ...
, 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

*''
Sedum ''Sedum'' is a large genus of flowering plants in the family Crassulaceae, members of which are commonly known as stonecrops. The genus has been described as containing up to 600 species, subsequently reduced to 400–500. They are leaf succule ...
'', a genus related to ''Dudleya''. *''
Echeveria ''Echeveria'' is a large genus of flowering plants in the family (biology), family Crassulaceae, native plant, native to semi-desert areas of Central America, Mexico and northwestern South America. Description Echeveria plants are evergreen. Flo ...
'', a genus similar to ''Dudleya''. *
Reid Moran Reid Venable Moran (June 30, 1916 – January 21, 2010) was an American botanist and the curator of botany at the San Diego Natural History Museum from 1957 to 1982. Moran was the world authority on the Crassulaceae, a family of succulent plants, ...
, who contributed greatly to the taxonomy of the genus. *
Nathaniel Lord Britton Nathaniel Lord Britton (1859 – 1934) was an American botanist and taxonomist who co-founded the New York Botanical Garden in the Bronx, New York (state), New York. Early life Britton was born on the 15 of January 1859 at New Dorp, Staten Island ...
and
Joseph Nelson Rose Joseph Nelson Rose (January 11, 1862 – May 4, 1928) was an American botanist. He was born in Union County, Indiana. His father died serving during the Civil War when Joseph Rose was a young boy. He later graduated from high school in Liberty, ...
, who described the genus.


References


Further reading

*Thompson, P. ''Dudleya and Hassenthaus Handbook''. Bonsall Publications. 1993.
Treatment from the Jepson Manual
{{Taxonbar, from1=Q138291, from2=Q87176181, from3=Q15733393 Crassulaceae genera Flora of the Southwestern United States Flora of Northwestern Mexico Succulent plants