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''Massalongiaceae'' is a small
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
in the order
Peltigerales Peltigerales is an order of lichen-forming fungi belonging to the class Lecanoromycetes in the division Ascomycota. The taxonomy of the group has seen numerous changes; it was formerly often treated as a suborder of the order Lecanorales. It con ...
. It has three genera and seven species. Species in this family have
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
as their primary symbiotic partner (
cyanobiont Cyanobionts are cyanobacteria that live in symbiosis with a wide range of organisms such as terrestrial or aquatic plants; as well as, algal and fungal species. They can reside within extracellular or intracellular structures of the host. In order ...
), so they belong to the group known as "cyanobacterial lichens".


Taxonomy

The family was
circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...
by Mats Wedin,
Per Magnus Jørgensen Per Magnus Jørgensen (born 1944) is a Norwegian botanist and lichenologist, and Professor Emeritus of systematic botany at the University of Bergen. He is known for his work on the lichen families Pannariaceae and Collemataceae. Jørgensen wa ...
, and Elisabeth Wiklund in 2007.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
analysis showed that the three genera formed a well-supported
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group that was distinct from other similar families in the Peltigerales. The family was named after the
type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...
, '' Massalongia''; the genus name honours Italian lichenologist
Abramo Bartolommeo Massalongo Abramo Bartolommeo Massalongo (13 May 1824 – 25 May 1860) was an Italian paleobotanist and lichenologist. He was born in Tregnago in the Province of Verona and took a great interest in botany as a young man. Massalongo joined the faculty of med ...
(1824-1860). Massalongiaceae has a
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
relationship with the family Peltigeraceae, and these two families together form a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
that is sister to a clade containing the families Vahiellaceae and Koerberiaceae. Previous phylogenetic studies had placed genus ''Massalongia'' in a group together with Peltigeraceae, Nephromataceae and Lobariaceae, although the relationships of ''Massalongia'' within this group were not precisely determined.


Description

The
thalli Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms were ...
of Massalongiaceae lichens range in form from
squamulose A squamulose lichen is a lichen that is composed of small, often overlapping "scales" called squamules. If they are raised from the substrate and appear leafy, the lichen may appear to be a foliose lichen, but the underside does not have a "skin ...
(scaley) to microfoliose (tiny and leafy) or microfruticose (tiny and bushy). The
photobiont A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
l genus ''
Nostoc ''Nostoc'', also known as star jelly, troll’s butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch’s jelly, is the most common genus of cyanobacteria found in vari ...
''. In the squamulose to microfoliose species, the lobes are rounded to elongated, and the branches in microfruticose species are terete (more or less circular in cross-section) and delicate. The
ascomata An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
are in the form of
apothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
, and they are usually biatorine (where the outer layers of the apothecium are light-coloured and contain no algae). The hamathecium (a collective term for all kinds of
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e or other tissues between asci) consists of unbranched to branched
paraphyses Paraphyses are erect sterile filament-like support structures occurring among the reproductive apparatuses of fungi, ferns, bryophytes and some thallophytes. The singular form of the word is paraphysis. In certain fungi, they are part of the fe ...
, and is
amyloid Amyloids are aggregates of proteins characterised by a Fibril, fibrillar morphology of 7–13 Nanometer, nm in diameter, a beta sheet (β-sheet) Secondary structure of proteins, secondary structure (known as cross-β) and ability to be Staining, ...
. The
asci ASCI or Asci may refer to: * Advertising Standards Council of India * Asci, the plural of ascus, in fungal anatomy * Accelerated Strategic Computing Initiative * American Society for Clinical Investigation * Argus Sour Crude Index * Association of ...
are semifissitunicate (i.e., discharge of spores from the ascus involves the separation of wall layers), with a thin apical tholus (the thickened inner part of an ascus tip) with an amyloid ring-structure, and cylindrical. The structure of the
asci ASCI or Asci may refer to: * Advertising Standards Council of India * Asci, the plural of ascus, in fungal anatomy * Accelerated Strategic Computing Initiative * American Society for Clinical Investigation * Argus Sour Crude Index * Association of ...
is particularly important in distinguishing between related families in the Peltigerales. For example, Peltigeraceae has a distinctive apical tube, while Nephromataceae lacks any amyloid apical structure, and
Lobariaceae The Peltigeraceae are a family of lichens in the order Peltigerales. The Peltigeraceae, which contains 15 genera and about 600 species, has recently (2018) been emended to include the families Lobariaceae and Nephromataceae. Many Peltigeraceae sp ...
has an indistinctive amyloid layer that is not comparable to the apical cap found in the asci of the Massalongiaceae. There are eight ascospores in each ascus. The spores have
septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five coun ...
that divide the spore transversely, and have a shape ranging from spindle-shaped (fusiform) to
ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a surface that may be defined as the ...
. The
conidiomata Conidiomata (singular: Conidioma) are blister-like fruiting structures produced by a specific type of fungus called a coelomycete. They are formed as a means of dispersing asexual spores call conidia, which they accomplish by creating the blister-li ...
are in the form of
pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...
; the
conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...
are
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 ...
, lack septa, and are rod-shaped to spindle-shaped. No
secondary compounds Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the norma ...
are known to occur in the family.


Habitat and distribution

Massalongiaceae lichens grow on rocks, and are often associated with
bryophyte The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in ...
s, growing among or on top of them. Collectively, the family has a
cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The ext ...
, but with more representation in northern
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
zones.


Genera and species

*'' Leptochidium'' ::*'' L. albociliatum'' ::*'' L. crenatulum'' *'' Massalongia'' ::*'' M. carnosa'' ::*'' M. griseolobulata'' ::*'' M. microphylliza'' ::*'' M. patagonica'' *'' Polychidium'' ::*'' P. muscicola'' Several additional species of ''Massalongia'' have been described, but some cannot be identified precisely due to lack of sufficient type material, as they are only known from the type collection, and some have been transferred to other genera.


References

{{Taxonbar, from=Q10577990 Peltigerales Lecanoromycetes families Taxa described in 2007 Taxa named by Per Magnus Jørgensen Lichen families