Hecastocleis shockleyi
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''Hecastocleis'' is a genus of low thorny shrubs with stiff branches, assigned to the daisy family. At the tip of each of the branches, inflorescences are subtended by oval, thorny, whitish to greenish bracts that enclose several flower heads which each contain only one pinkish bud, opening into a white corolla. It contains but one species, ''Hecastocleis shockleyi'', the only representative of the tribe Hecastocleideae, and of the subfamily Hecastocleidoideae. Its vernacular name is prickleleaf. It is confined to the southwestern United States. and


Description

''Hecastocleis shockleyi'' is a
xerophytic A xerophyte (from Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or places in the Alps or t ...
thorny shrub of , occasionally 1½ m (4 ft 11 in) high. It has sixteen chromosomes (2n=16).


Leaves

The leaves are alternately set along the branches. The leaf blades are hairless or with a few soft hairs, slightly olive green, stiff and leathery, with three main veins,
linear Linearity is the property of a mathematical relationship ('' function'') that can be graphically represented as a straight line. Linearity is closely related to '' proportionality''. Examples in physics include rectilinear motion, the linear ...
to narrowly ovate in shape, long, their base approximately clasping the branch, with thickened margins usually carrying three to six spines of long, denser spaced near the base, while the tip is more or less abruptly tapering into a point.


Inflorescence

The complex inflorescences are carried at the end of the branches. These consist of a number of crowded clusters. Each of the clusters is subtended by white to yellowish green, wavy, ovate to orbicular bracts that have a spiny margin, and further consist of one to five flower heads which each contain only a single
disk floret The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae we ...
. The most outward part of the flower head is the
involucre In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or o ...
, which is narrowly vase-shaped to cylindric and approximately high, and consists of about six worls of four bracts called
phyllaries In botanical terminology, a phyllary, also known an involucral bract or tegule, is a single bract of the involucre of a composite flower. The involucre is the grouping of bracts together. Phyllaries are reduced leaf-like structures that form one o ...
, which have often soft woolly hairs around the edge. The base of the flower head (or receptacle) is flat and lacks a bract directly at the base of the floret. The corolla of the florets is pinkish purple when still in bud, but turns pinkish white at flowering, at which time it is about high. These are hermaphrodite, star-symmetric (or
actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
), and have five narrow outwards oriented lobes. The five pinkish purple anthers are fused into a long tube, that initially covers the entire style. The bone-colored style grows through the anther tube, collecting the pollen on hairs, and displaying it above the anter tube like in all other Asteraceae. The style branches are only and have rounded tips. At the base of the corolla the one-seeded indehiscent fruit (called cypselas) develops, which is not flattened, has four to five hardly visible nerves, turns brown and has lost any hairs when ripe, and is about ½ cm (0.2 in) high. On the top of the cypselas is a crown of six unequal scales (the pappus) of high. These scales are lanceolate in shape, may have several teeth and are sometimes fused at their base, so forming a crown. The pollen of ''Hecastocleis'' is yellow, unadorned and tricolpate.


Differences with other Asteraceae

Like ''Hecastocleis'', some other Asteraceae also have flower heads consisting of a single floret, such as ''
Gundelia ''Gundelia'' is a low to high (20–100 cm) thistle-like perennial herbaceous plant with latex, spiny compound inflorescences, reminiscent of teasles and eryngos, that contain cream, yellow, greenish, pink, purple or redish-purple disk flo ...
'', a perennial herbaceous plant from the Middle-East, and '' Gymnarrhena'' a winter annual from northern Africa and the Middle-East. Both have male flowers and female flowers, not hermaphrodite as in ''Hecastocleis'', while ''Gymnarrhena'' has (trimerous or) tetramerous male florets, not pentamerous.


Taxonomy

''Hecastocleis shockleyi'' was described by Asa Gray in 1882 and placed it to the tribe
Mutisieae Mutisieae is a tribe of the family Asteraceae, subfamily Mutisioideae. Genera Mutisieae genera recognized by the Global Compositae Database as of June 2022: *'' Adenocaulon'' *'' Amblysperma'' *'' Brachyclados'' *''Chaetanthera'' *'' Chap ...
.
Kåre Bremer Kåre Bremer (born 17 January 1948) is a Swedish botanist and academic. He has also been Vice-Chancellor of Stockholm University. Career Professor Bremer received his doctorate in Botany from Stockholm University in 1976, where he worked as ...
in 1994 assigned it to the subtribe Mutisiinae, while Hind in 2006 erected its own group within the Mutisieae.


Modern classification

Scholars agree that ''Hecastocleis'' occupies a very isolated position on the
evolutionary grade A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of sp ...
of subfamilies of the Asteraceae, and this is expressed by the erection of the monotypic subfamily Hecastocleidoideae.


Phylogeny

The current insights in the relationships between ''Hecastocleis'' and its closest relatives, as based on genetic analysis, is expressed in the following tree.


Biogeology

Earlier splits of the Asteraceae tree are supposed to have taken place in southern South America. The splits that occur later than ''Hecastocleis'' have occurred Africa, especially southern Africa, and some may also have occurred in Asia. This implies that the common ancestor of ''Hecastocleis'' and all later branches moved from South America to North American and after the split with ''Hecastocleis'', the ancestor of the remainder of the subfamilies arrived in Africa from North-America. Alternatively, only the ancestor of ''Hecastocleis'' migrated from South America to North America while the ancestor of the remainder was distributed from South America to Africa. The placement of ''Hecastocleis'' in the tree could also be incorrect due to parallel evolution an reversal of earlier mutations. However, the support in the analysis for the separation of ''Hecastocleis'' from the Gochnatieae is strong.


Etymology

The name of the genus ''Hecastocleis'' is the contraction of two Greek words, ''ἕκαστος'' (hekastos) meaning "each" and ''κλειω'' (kleio) which is said to mean "to shut up", referring to each flower having its own involucre. An alternative meaning of ''kleio'' is "glory". The species was named ''shockleyi'' in honor of William Hillman Shockley, an early plant collector of the flora of Nevada and father of the co-inventor of the transistor
William Shockley William Bradford Shockley Jr. (February 13, 1910 – August 12, 1989) was an American physicist and inventor. He was the manager of a research group at Bell Labs that included John Bardeen and Walter Brattain. The three scientists were jointl ...
.


Distribution

Prickleleaf is native to the desert plains and mountains of eastern California ( Inyo,
Mono Mono may refer to: Common meanings * Infectious mononucleosis, "the kissing disease" * Monaural, monophonic sound reproduction, often shortened to mono * Mono-, a numerical prefix representing anything single Music Performers * Mono (Japanese b ...
,
Kern KERN (1180 AM broadcasting, AM) is a commercial radio, commercial radio station city of license, licensed to Wasco, California, Wasco-Greenacres, California, and serving the Bakersfield metropolitan area. The station is owned by American General ...
, and San Bernardino Counties) and southern Nevada (
Mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
, Esmeralda, Nye,
Lincoln Lincoln most commonly refers to: * Abraham Lincoln (1809–1865), the sixteenth president of the United States * Lincoln, England, cathedral city and county town of Lincolnshire, England * Lincoln, Nebraska, the capital of Nebraska, U.S. * Lincol ...
, and Clark Counties), where it grows on arid, rocky slopes and flats.Flora of North America, Vol. 19, 20 and 21 Page 71, ''Hecastocleis'' A. Gray
/ref> Records of this plant include
Mount Charleston Mount Charleston, including Charleston Peak (Nuvagantu, literally "where snow sits", in Southern Paiute or Nüpakatütün in Shoshoni) at , is the highest mountain in both the Spring Mountains and Clark County, in Nevada, United States. It is ...
,
Nevada Test Site The Nevada National Security Site (N2S2 or NNSS), known as the Nevada Test Site (NTS) until 2010, is a United States Department of Energy (DOE) reservation located in southeastern Nye County, Nevada, about 65 miles (105 km) northwest of the ...
,
Grapevine Mountains The Grapevine Mountains are a mountain range located along the border of Inyo County, California and Nye County, Nevada in the United States. The mountain range is about long and lies in a northwest-southeasterly direction along the Nevada-Califor ...
and
Red Pass Red Pass is a gap in the Avawatz Mountains, in San Bernardino County, California. Red Pass, lies between the Silurian Valley and the valley drained by an as yet unnamed tributary of Salt Creek, which drains much of the area of Fort Irwin Nation ...
.


Ecology

In the mountains rimming the north-side of Death Valley, ''Hecastocleis shockleyi'' may be one of the dominant shrubs together with ''
Atriplex confertifolia ''Atriplex confertifolia'', the shadscale or spiny saltbush, is a species of evergreen shrub in the family Amaranthaceae, which is native to the western United States and northern Mexico. Description The height of ''Atriplex confertifolia'' ...
'', ''
Eriogonum fasciculatum ''Eriogonum fasciculatum'' is a species of wild buckwheat known by the common names California buckwheat and flat-topped buckwheat. Characterized by small, white and pink flower clusters that give off a cottony effect, this species grows vari ...
'', and ''
Tetradymia axillaris ''Tetradymia axillaris'' is a flowering plant in the daisy family known by the common names longspine horsebrush and cottonthorn. This is a plant of the sagebrush and desert plant communities of the southwestern United States. The plant forms a ...
''. Less dominant but often present shrubs are ''
Ephedra viridis ''Ephedra viridis'', known by the common names green Mormon tea, green ephedra, and Indian tea, is a species of ''Ephedra''. It is indigenous to the Western United States, where it is a member of varied scrub, woodland, desert, and open habitats. ...
'', ''
Ericameria laricifolia ''Ericameria laricifolia'' is a North American species of flowering shrub in the family Asteraceae known by the common name turpentine bush, or turpentine-brush. It is native to the southwestern United States (Arizona, New Mexico, western Texas ...
'', and '' Lepidium fremontii''. Trees are absent. Herbaceous plants form an open layer and regularly include the grasses '' Achnatherum hymenoides'', '' Bromus rubens'' and ''
Poa secunda ''Poa secunda'' (variously known by the common names of Sandberg bluegrass, alkali bluegrass, big bluegrass, Canby's bluegrass, Nevada bluegrass, one-sided bluegrass, Pacific bluegrass, pine blugrass, slender bluegrass, wild bluegrass, and curly ...
'', in addition to ''
Claytonia perfoliata ''Claytonia perfoliata'' ( syn. ''Montia perfoliata''), also known as miner's lettuce, Indian lettuce, winter purslane, or ''palsingat'' (Cahuilla), is a flowering plant in the family Montiaceae. It is an edible, fleshy, herbaceous, annual plant ...
'', '' Cryptantha utahensis'', '' Delphinium parishii, several '' Gilia species, and '' Phacelia vallis-mortae''.
Biological soil crust Biological soil crusts are communities of living organisms on the soil surface in arid and semi-arid ecosystems. They are found throughout the world with varying species composition and cover depending on topography, soil characteristics, climate, ...
, lichen, and moss can also be found in this plant community. It is found primarily on steep, very rocky slopes and bedrock outcrops with northerly aspects. ''Hecastocleis shockleyi'' occurs at altitudes between 1250 and 1600 m. It typically grows on calcareous clay loam, loamy sand, or sandy clay that has resulted from erosion of dolomite, limestone or shale, and is poor in nutrients. Pollination of ''Hecastocleis'' has not yet been observed, but as its flowers are white, nocturnal insects are likely candidates.


References


External links


Jepson Manual TreatmentUSDA Plants ProfilePhoto gallery
{{Taxonbar, from1=Q16056057, from2=Q142364, from3=Q16056058, from4=Q5893343 Asteraceae Monotypic Asteraceae genera Flora of Nevada Flora of the California desert regions Plants described in 1882 Flora without expected TNC conservation status