Ptilidium Pulcherrimum Sporen
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''Ptilidium'' is a genus of
liverwort Liverworts are a group of non-vascular land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry ...
, and is the only
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 ...
in
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Ptilidiaceae. It includes only three species: '' Ptilidium californicum'', '' Ptilidium ciliare'', and '' Ptilidium pulcherrimum''. The genus is distributed throughout the arctic and subarctic, with disjunct populations in New Zealand and Tierra del Fuego. Molecular analysis suggests that the genus has few close relatives and diverged from other leafy liverworts early in their evolution.


Description

The name of the genus comes from the Greek word ''ptilidion'' for "small feather", in reference to the multiply deeply divided leaves with fringed edges, which give the plant a "feathery" appearance. Unlike other
leafy liverwort Jungermanniales is the largest Order (biology), order of Marchantiophyta, liverworts. They are distinctive among the liverworts for having thin leaf-like flaps on either side of the stem. Most other liverworts are thalloid, with no leaves. Due to ...
s, the underleaves are not significantly smaller than the lateral leaves. The "flossy" appearance from the leaf edges, together with the characteristic yellowish-brown or reddish-brown color make the genus easy to recognize. Like ''Ptilidium'', '' Blepharostoma'' and '' Trichocolea'' have deeply divided leaves with marginal cilia, however ''Ptilidium'' differs from these other two genera in that its leaf cells have bulging trigones (thickenings at the corners between
cell wall A cell wall is a structural layer that surrounds some Cell type, cell types, found immediately outside the cell membrane. It can be tough, flexible, and sometimes rigid. Primarily, it provides the cell with structural support, shape, protection, ...
s). The plants grow in dense mats, with stems growing either prostrate or ascending. Individual stems are once or twice
pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and ...
, rarely with branches and only a few short
rhizoid Rhizoids are protuberances that extend from the lower epidermal cells of bryophytes and algae. They are similar in structure and function to the root hairs of vascular land plants. Similar structures are formed by some fungi. Rhizoids may be un ...
s. The leaves are incubous and divided deeply into three to five portions, and edges of the leaf divisions are fringed with cilia. The underleaves are similar to the lateral leaves, but are slightly smaller. All species are
dioicous Dioicy () is a sexual system in non-vascular plants where archegonia (female organs) and antheridia (male organs) are produced on separate plants in the gametophyte phase. It is one of the two main sexual systems in bryophytes, the other being ...
, producing
antheridia An antheridium is a haploid structure or organ producing and containing male gametes (called ''antherozoids'' or sperm). The plural form is antheridia, and a structure containing one or more antheridia is called an androecium. The androecium is al ...
and
archegonia An archegonium (: archegonia), from the Ancient Greek ''ἀρχή'' ("beginning") and ''γόνος'' ("offspring"), is a multicellular structure or organ of the gametophyte phase of certain plants, producing and containing the ovum or female gamet ...
on separate plants. The archegonia are terminal on a main stem. Mature
sporophyte A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
s develop from within a large perianth with three distal folds. The three species in the genus may be distinguished by the density of cilia along the leaf margin, depth of lobing in the leaf, width of the leaf base, and the substrate on which it is found growing. ''P. californicum'' has few cilia along the edge of its leaves and has underleaves lobed to about seven-eighths of their length. The other two species have many marginal cilia and underleaves lobed to no more than half their length. ''P. ciliare'' usually grows on soil and has leaf lobes that are 15–20 cells wide at their base. ''P. pulcherrimum'' usually grows on wood or rock, and has leaf lobes normally 6–10 cells wide at their base.


Distribution

The genus ''Ptilidium'' has a
boreal Boreal, northern, of the north. Derived from the name of the god of the north wind from Ancient Greek civilisation, Boreas (god), Boreas. It may also refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernand ...
distribution, and is found in abundance in
coniferous forest Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
s of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, and
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 ...
, as well as in
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
and
Tierra del Fuego Tierra del Fuego (, ; Spanish for "Land of Fire", rarely also Fireland in English) is an archipelago off the southernmost tip of the South America, South American mainland, across the Strait of Magellan. The archipelago consists of the main is ...
. Plants often grow attached to the bark of trees in the northern hemisphere, but may occur in rocks in mountain districts of New Zealand. At the more temperate ends of its range, plants are restricted to higher elevations. Schuster (1984) proposed that the disjunct distribution of '' Ptilidium ciliare'' between the northern and southern hemispheres could be explained by migration of the
Indian Plate The Indian plate (or India plate) is or was a minor tectonic plate straddling the equator in the Eastern Hemisphere. Originally a part of the ancient continent of Gondwana, the Indian plate broke away from the other fragments of Gondwana an ...
from
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
. In this hypothesis, ''P. ciliare'' is a species originally native to Gondwana, and sterile populations existing in modern New Zealand and Tierra del Fuego are
relict A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
s of this earlier distribution. The other two species of ''Ptilidium'' are thus believed to be later descendants. Schuster's hypothesis is partially based upon a belief that ''Ptilidium'' is related to the genera ''
Mastigophora A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
'' and '' Dendromastigophora'', both of which are largely restricted to the southern hemisphere. However, this relationship is not supported by modern molecular analysis, which places ''Mastigophora'' in an entirely different part of the liverwort phylogeny. Instead, ''Ptilidium'' is now believed to be part of an isolated clade allied only to two East Asian endemics, and it is thus more likely that the sterile populations of ''Ptilidium'' in the southern hemisphere reflect long-distance dispersal of plant fragments. ''Ptilidium ciliare'' is tolerant of desiccation and is ubiquitous in the Arctic, but rarely produces
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s, and it is therefore believed to spread by means of such fragments.


Phylogeny

The diagram at left summarizes a portion of a 2006
cladistic analysis Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
of liverworts based upon three chloroplast genes, one nuclear gene, and one mitochondrial gene. The genus '' Trichocoleopsis'' was not included in the original broad analysis, but is the sister taxon of '' Neotrichocolea'' according to a more narrowly focussed study utilizing six chloroplast genes, two nuclear genes, and a mitochondrial gene. The genus ''Ptilidium'' is sister to the ''Trichocoleopsis''-''Neotrichocolea'' clade. This combined clade, in turn, attaches at the base of a large clade (2600 species) designated "Leafy II". That clade, together with "Leafy I" (another 1800 species) and '' Pleurozia'' constitute the
Jungermanniales Jungermanniales is the largest Order (biology), order of Marchantiophyta, liverworts. They are distinctive among the liverworts for having thin leaf-like flaps on either side of the stem. Most other liverworts are thalloid, with no leaves. Due to ...
, as traditionally defined. ''Ptilidium'', ''Neotrichocolea'', and ''Trichocoleopsis'' thus sit at the base of the Jungermanniales, at a point where the two major groups of leafy liverworts diverge from each other.


Gallery

File:Ptilidium ciliare blatt.jpeg , File:Ptilidium ciliare laminazellen.jpeg ,


References


External links

* * {{Taxonbar, from1=Q310414, from2=Q16916366 Ptilidiales Liverwort genera