Harfordia Macroptera
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''Harfordia macroptera'' is a woody
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
in the family
Polygonaceae The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus ''Polygonum'', and was first used by Antoine Laurent de Jussieu in 1789 ...
commonly known as rabbit's purse. It is the sole species in the genus ''Harfordia'', which is
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to the Baja California Peninsula. This plant is characterized by a unique, conspicuous bladder embellished with red to purple veins that surround the fruit. The sac-like structure is actually a dramatically modified
bract 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 of ...
, and assists in dispersal of the seeds through the wind.


Description

This sprawling to spreading
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
or
subshrub A subshrub (Latin ''suffrutex'') or dwarf shrub is a short shrub, and is a woody plant. Prostrate shrub is a related term. "Subshrub" is often used interchangeably with "bush".Jackson, Benjamin, Daydon; A Glossary of Botanic Terms with their Der ...
is characterized by its conspicuous and colorful sac-like
bract 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 of ...
s that aid in dispersal of the
achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not ope ...
. Plants may be
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...
but are more commonly
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
. This species may not be immediately obvious in habitat until their inflated bracts are fully expanded.


Morphology

This plant has numerous slender to stout stems that arise from a woody
taproot A taproot is a large, central, and dominant root from which other roots sprout laterally. Typically a taproot is somewhat straight and very thick, is tapering in shape, and grows directly downward. In some plants, such as the carrot, the taproo ...
. The leaves are cauline, opposite, with the leaf blades shaped linear to oblanceolate to obovate-spatulate. The leaves may be glabrous or
pubescent The adjective pubescent may describe: * people or animals undergoing puberty * plants that are hairy, covered in trichomes * insects that are covered in setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a ...
with slightly curled hairs. The leaves are thin to thick-ish and more or less
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 ...
. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
s have flowers borne in axillary clusters, or on perfectA "perfect" flower contains both male and female organs. or female plants, on short, stout peduncles. The peduncles are 0.7 to 2 cm long, and are usually densely pubescent. The involucres are lacking in male plants, and in perfect or female plants the involucre is reduced to a highly modified bract. This highly modified bract is a sac or bladder-like structure that enlarges with development of the flower, colored
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
and textured with deep red to red-purple veins, and lacking hairs. The flowers are either imperfectAn "imperfect" flower contains either male or female organs. or perfect, with the 6
tepal A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...
s monomorphic and shaped lanceolate. The male flowers are pale yellow, smaller than the female ones, and have an occasionally have an aborted
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
. The female flowers are initially pale yellow but become red to rose, and always have aborted
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s. The perfect flowers are pale yellow to rose. There are 9 stamens, with those on the male flowers slightly protruding on elongated filaments that bear yellow oval anthers, while those on the female flower on minute filaments bearing small aborted anthers. The
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
is an
achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not ope ...
, 3 to 5 mm long, colored yellowish-brown to brown.


Taxonomy


Classification

The first plants discovered were initially described as being in the genus '' Pterostegia''. However, upon realization that ''Pterostegia macroptera'' was a woody
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
, unlike the type species ''Pterostegia drymarioides'' (which was an
annual Annual may refer to: * Annual publication, periodical publications appearing regularly once per year **Yearbook ** Literary annual * Annual plant * Annual report * Annual giving * Annual, Morocco, a settlement in northeastern Morocco * Annuals (b ...
), and that the large, inflated
bract 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 of ...
s were unique,
Edward Lee Greene Edward Lee Greene (August 20, 1843–November 10, 1915) was an American botanist known for his numerous publications including the two-part ''Landmarks of Botanical History'' and the describing of over 4,400 species of plants in the American W ...
and Charles C. Parry established the genus ''Harfordia'', placing within it ''Harfordia macroptera'', ''Harfordia galioides'' and the
Cedros Island Cedros Island (''Isla de Cedros'', "island of cedars" in 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 of which includes ...
endemic ''Harfordia fruticosa''.
Ira Loren Wiggins Ira Loren Wiggins (1 January 1899 – 28 November 1987) was an American botanist, Curator of the Dudley Herbarium, and Director of the Natural History Museum (1940–1962) at Stanford University. He was a Stanford faculty member from 1929 until h ...
only accounted for ''H. macroptera'' in his flora of the
Sonoran Desert The Sonoran Desert ( es, Desierto de Sonora) is a desert in North America and ecoregion that covers the northwestern Mexican states of Sonora, Baja California, and Baja California Sur, as well as part of the southwestern United States (in Arizona ...
, citing ''Pterostegia galioides'' as a synonym and disposing of ''H. fruticosa'' due to his exclusion of Cedros Island endemics.
James L. Reveal James Lauritz Reveal (March 29, 1941 – January 9, 2015) was a U.S. botanist best known for his contributions to the genus '' Eriogonum'' and for his work on suprageneric names. His website, at PlantSystematics.org, also presents material on plan ...
took into account the three distinct entities Greene had described (''galioides, fruticosa'' and ''macroptera''), and opted to place them as varieties within a
monospecific In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
genus. His basis for separation was the fact that the varieties were primarily
allopatric Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
, only overlapping in one small area in northwestern Baja California Sur. Where the plants overlap, a series of morphological intermediates between them are found.


Varieties

Although the three varieties are weakly defined morphologically, they are geographically distinct. They are: * ''Harfordia macroptera'' var. ''galioides'' (Vizcaino rabbit's purse) – A large and often sprawling shrub that is 30 to 60 cm tall and 40 to 80 cm across. The leaves are shaped linear to narrowly oblanceolate, and are 0.8 to 2 cm long, and 0.5 to 2 mm wide, hairy with slightly curled hairs. The sac-like bract is 2 to 3 cm across at maturity. Chromosome count is ''n'' = 20. This variety is uncommonly distributed from San Vicente in northwestern Baja California to the Vizcaino Peninsula of northern
Baja California Sur Baja California Sur (; 'South Lower California'), officially the Free and Sovereign State of Baja California Sur ( es, Estado Libre y Soberano de Baja California Sur), is the least populated state and the 31st admitted state of the 32 federal ent ...
. * ''Harfordia macroptera'' var. ''fruticosa'' (Cedros rabbit's purse) ''–'' A large sprawling shrub that is 10 to 30 cm tall and 20 to 40 cm across. The leaves are broadly oblanceolate to obovate-spatulate, and are 0.2 to 0.8 cm long, and 2 to 4 mm wide, lacking hairs and thick and succulent. The sac-like bract is 1 to 2 cm across at maturity. This species is endemic to
Cedros Island Cedros Island (''Isla de Cedros'', "island of cedars" in 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 of which includes ...
off of the coast of Baja California. * ''Harfordia macroptera'' var. ''macroptera'' (Magdalena rabbit's purse) – A large sprawling shrub that is 20 to 40 cm tall and 40 to 60 cm across. The leaves are broadly oblanceolate to spatulate, and 0.5 to 1.5 cm long, 2 to 6 mm wide, lacking hairs, and thick and succulent. The sac-like bract is 1 to 2 cm across at maturity. This species is common on gravelly soil in the
Magdalena Bay Magdalena Bay ( es, Bahía Magdalena) is a long bay in Comondú Municipality along the western coast of the Mexican state of Baja California Sur. It is protected from the Pacific Ocean by the unpopulated sandy barrier islands of Isla Magdalen ...
region and surrounding islands in Baja California Sur.


Phenology

The characteristic bladder or sac-like structure that surrounds the
achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not ope ...
originally starts as a tiny immature
bract 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 of ...
subtending the
pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
. The bract is initially reflexed and composed of three individual segments that only join each other at the point of attachment. As the bract matures, it enlarges and the forms a sheath around the pedicel. As soon as the flower inside is
fertilized Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
, the bract rapidly expands upward and outwards. Eventually, the whole structure completely overtops and encompasses the achene. The two laminar sides of the bract continue to expand to form the conspicuous bladder-like portion of the bract. The modified bract is hypothesized to aid in the dispersal of the achene. The inflated and papery bract is easily detached and blow through the wind, with the achene inside. This leaves the fully mature achene within able to be readily transported. The bract does dry, but as it is filled with veins, it does not simply disintegrate. Even after detachment, rain can re-inflate and move with the help of the winds associated with thunderstorms. The achene can remain with the bracts for up to two to three years.


Notes


References


External links


U.C. Photo Gallery — ''Harfordia macroptera''
{{Taxonbar, from1=Q10293910, from2=Q15928023 Polygonaceae Flora of Baja California Flora of Baja California Sur Endemic flora of Mexico