Riella Echinospora
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''Riella'' is a genus in the
liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
family
Riellaceae Sphaerocarpales is an order of plants within the liverworts. Approximately twenty species are in this order which is sub-divided into four families: Monocarpaceae, Sphaerocarpaceae and Riellaceae, as well as the extinct family Naiaditaceae. T ...
, and includes about eighteen species. Plants in the genus are small and grow submerged in shallow temporary pools. Although the genus is widely distributed in the Northern Hemisphere, locating populations is often difficult. Its occurrence is sporadic and local, and the tiny plants are ephemeral. The ornamented
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s remain viable for several years, allowing the plants to survive annual drying of their habitat. The plants are easily grown in laboratory cultures.


Description

''Riella'' plants are small, usually or less, and
thalloid Thallus (plural: thalli), from Latinized Ancient Greek, Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some Marchantiophyta, liverworts, lichens, and the Myxogastr ...
, with an appearance like an immature
alga Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mi ...
. The plant consists of an erect central axis ("stem") that is commonly forked, but only sparingly, and the plants are bright green. The stem bears a thin dorsal
lamina Lamina may refer to: Science and technology * Planar lamina, a two-dimensional planar closed surface with mass and density, in mathematics * Laminar flow, (or streamline flow) occurs when a fluid flows in parallel layers, with no disruption betwee ...
or "wing", which being on only one side gives the plant an asymmetrical appearance. This lamina is ruffled or undulate. Early descriptions of the species ''
Riella helicophylla ''Riella'' is a genus in the liverwort family Riellaceae, and includes about eighteen species. Plants in the genus are small and grow submerged in shallow temporary pools. Although the genus is widely distributed in the Northern Hemisphere, loca ...
'' overemphasized the spiralled form of the lamina, which does not occur in other species. One species, the Algerian '' Riella bialata'' bears two wings along its stems instead of the usual single lamina. The lamina has a thickness of only a single layer of cells, with all the cells thin- walled and chlorophyllose. In addition to the lamina, the central stem bears delicate, small, leaf-like scales in three series along its lateral and ventral aspects. The leaf scales are dimorphic and contain scattered oil cells, with a single
oil body An oil body is a lipid-containing structure found in plant cells. The term can refer to at least two distinct kinds of structures in different kinds of plants. Oil bodies in liverworts Liverwort complex oil bodies are structures unique to live ...
per cell. ''Riella'' is thus the only member of its order with oil cells, although other genera in the class
Marchantiopsida Marchantiopsida is a class of liverworts within the phylum Marchantiophyta. The species in this class are known as complex thalloid liverworts. The species in this class are widely distributed and can be found worldwide. Phylogeny Based on the w ...
commonly have them.


Distribution

Although found nearly worldwide, ''Riella'' populations are sporadic and limited to only a few known localities. The plants are never common, and are found most often in places where the summer is dry and the winter is wet and mild. Prior to 1900, ''Riella'' was thought to occur only in regions around the
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
. The first collections from outside this region were published in 1902 when Porsild reported collecting ''R. paulsenii'' in
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a transcontinental country located mainly in Central Asia and partly in Eastern Europe. It borders Russia to the north and west, China to the east, Kyrgyzstan to the southeast, Uzbeki ...
in 1898. Three further specimens from the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
were reported in 1903, although one of these had been collected much earlier in 1855. Since that time, additional populations have been reported from
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
, southern
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
, and southeastern
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
. ''Riella'' species primarily grow as submerged
aquatic plant Aquatic plants are plants that have adapted to living in aquatic environments (saltwater or freshwater). They are also referred to as hydrophytes or macrophytes to distinguish them from algae and other microphytes. A macrophyte is a plant that ...
s, rarely floating or emergent. Although it is unique among hepatics as a submerged aquatic, it is not the only aquatic liverwort. The plants grow in fresh or brackish water of temporary bodies of water, and rarely in permanent ones.


Classification

The genus ''Riella'' was first published in 1852 by
Camille Montagne Jean Pierre François Camille Montagne (15 February 1784 – 5 December 1866) was a French military physician and botanist who specialized in the fields of bryology and mycology. He was born in the commune of Vaudoy in the department of Seine-et- ...
. The group was given its own family Riellaceae 40 years later by
Adolf Engler Heinrich Gustav Adolf Engler (25 March 1844 – 10 October 1930) was a German botanist. He is notable for his work on alpha taxonomy, plant taxonomy and phytogeography, such as ''Die natürlichen Pflanzenfamilien'' (''The Natural Plant Families'' ...
. Modern bryologists classify the Riellaceae in the order Sphaerocarpales, along with the family
Sphaerocarpaceae Sphaerocarpaceae is a family of liverworts known as bottle liverworts. Approximately ten species are included in this family, most of them in the genus ''Sphaerocarpos ''Sphaerocarpos'' is a genus of plants known as bottle liverworts. There ...
. Studies comparing genetic sequences of ''Riella'' plants support this relationship. Porsild recognized a subgenus ''Trabutiella'' distinguished by the presence of longitudinal wings on the involucre. The only other subgenus (''Riella'' subg. ''Riella'') bears smooth involucres that lack these wings of tissue. Species of ''Riella'' are distinguished using morphology of the spores or archegonial involucres, since the gametophytes of the genus possess few distinguishing non-reproductive characters. There are 19, 18, or 17 species in the genus.


References


External links

* *
"Riellaceae" in ''Bryophyta Flora of North America''
{{Taxonbar, from=Q7333114 Sphaerocarpales Liverwort genera Taxa named by Camille Montagne