Enigmotrema
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''Enigmotrema'' is a fungal
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 the family
Graphidaceae The Graphidaceae are a family of lichen-forming fungi in the order Graphidales. The family contains nearly a hundred genera and more than 2000 species. Although the family has a cosmopolitan distribution, most Graphidaceae species occur in tropic ...
. It contains only one known species, ''Enigmotrema rubrum'', a lichen which was discovered in Costa Rica. Both the genus and species were first described in 2012 by the lichenologist
Robert Lücking Robert Lücking (born 1964) is a German lichenologist, known for his extensive research on foliicolous lichens (lichens that live on leaves) and his significant contributions to the taxonomy, ecology, and biodiversity of fungi and lichens. He e ...
. The genus name combines the Greek words for "mystery" and "hole", referring to the unusual appearance of its reproductive structures. ''Enigmotrema'' is characterised by its olive-coloured body (
thallus Thallus (: 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. A thallus usually names the entir ...
) and distinctive dark red fruiting bodies (
apothecia An ascocarp, or ascoma (: ascomata), 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. As ...
) that undergo a unique opening process as they mature. The genus is closely related to '' Cruentotrema'' but differs in its genetic makeup and the development of its fruiting bodies. Currently, ''Enigmotrema'' is only known from a single location in a
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
rainforest in Costa Rica, where it was found growing on tree bark.


Taxonomy

The genus ''Enigmotrema'' was
circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...
by the lichenologist
Robert Lücking Robert Lücking (born 1964) is a German lichenologist, known for his extensive research on foliicolous lichens (lichens that live on leaves) and his significant contributions to the taxonomy, ecology, and biodiversity of fungi and lichens. He e ...
in 2012. The name ''Enigmotrema'' combines the Greek words ''enigma'' (meaning mystery or puzzle) and ''trema'' (meaning hole or pore), referring to the unusual appearance of its fruiting bodies. ''Enigmotrema'' is closely related to the genus '' Cruentotrema'' but differs in several key aspects. Genetic analysis of the mitochondrial small subunit ribosomal DNA (mtSSU) revealed unique characteristics that place ''Enigmotrema'' outside the ''Cruentotrema''-''Dyplolabia''
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, a group of closely related lichen genera. Later molecular analysis suggests that the species is nested with ''
Fissurina ''Fissurina'' is a genus of lichenized fungi in the family Graphidaceae. It has about 160 species, most of which are found in tropical regions. Taxonomy The genus was circumscription (taxonomy), circumscribed by the French botanist Antoine Lau ...
'', and that it is closely related to the '' Fissurina pseudostromatica'' clade.


Description

''Enigmotrema'' lichens possess a distinctive appearance that sets them apart from their close relatives. The
thallus Thallus (: 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. A thallus usually names the entir ...
of the lichen has an olive-green to olive-yellow colouration. Its surface is smooth to slightly uneven, protected by a dense outer layer, the . This cortex is composed of tightly interwoven fungal threads, or
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 o ...
e, forming a protective barrier for the lichen. A characteristic feature of ''Enigmotrema'' is its
apothecia An ascocarp, or ascoma (: ascomata), 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. As ...
. These fruiting bodies undergo a unique development process that gives the genus its enigmatic character. Initially, the apothecia appear as small, closed, dark red structures reminiscent of tiny flasks. As they mature, these structures, measuring between 0.7 and 1.2 millimetres in diameter, begin a transformation. First, the outer layer of the apothecium splits open, revealing a dark red inner layer. This is followed by the remaining covering splitting into triangular teeth-like structures that curl inwards. This process gradually exposes the brown-black surface, known as the , where spores are produced. The end result is a complex, open structure that bears little resemblance to its initial closed form. Inside these apothecia, ''Enigmotrema'' produces colourless,
ellipsoid An ellipsoid is a surface that can be obtained from a sphere by deforming it by means of directional Scaling (geometry), scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a Surface (mathemat ...
spores called . These spores are relatively small, measuring 10–12
micrometre The micrometre (English in the Commonwealth of Nations, Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a uni ...
s (μm) in length and 5–6 μm in width. They possess three cross-walls, or
septa SEPTA, the Southeastern Pennsylvania Transportation Authority, is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly four million people througho ...
, and have a distinctive internal structure characterised by thick walls and diamond-shaped chambers (. This internal arrangement is similar to that found in some other lichen genera, particularly ''
Astrothelium ''Astrothelium'' is a large genus of corticolous (bark-dwelling) lichens in the family Trypetheliaceae. The genus is characterized by a corticate thallus and diverse ascomata structures, which can be simple, aggregated, or forming . ''Astrotheli ...
''. From a chemical perspective, ''Enigmotrema'' is notable for containing a dark red
pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
called isohypocrellin in the inner layer (
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
) of its apothecia. This pigment contributes to the striking red colour of the fruiting bodies. Standard analytical techniques did not detect any other significant chemical substances in the lichen's composition.


Habitat and distribution

''Enigmotrema rubrum'', was discovered growing on tree bark in Costa Rica. The specimen was collected from the
Las Cruces Biological Station The Las Cruces Biological Station / Wilson Botanical Garden is located in the southern Puntarenas province of Costa Rica, and is the newest of the three research stations operated by the Organization for Tropical Studies (OTS). Las Cruces includ ...
at the
Las Cruces Biological Station The Las Cruces Biological Station / Wilson Botanical Garden is located in the southern Puntarenas province of Costa Rica, and is the newest of the three research stations operated by the Organization for Tropical Studies (OTS). Las Cruces includ ...
, located in the Talamanca mountain range. This area is characterised as a
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
rainforest zone at an elevation of about 1,200 metres above sea level. The lichen was found growing on the lower trunk of a tree in a semi-exposed location. While the immediate surroundings were described as "anthropogenic vegetation" (vegetation influenced by human activity), it was within the context of a protected
botanical garden A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens. is ...
.


References

{{Taxonbar, from1=Q107289178 , from2=Q107656257 Graphidaceae Taxa described in 2012 Graphidales genera Lichen genera Taxa named by Robert Lücking