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A herbarium specimen of the lichen Leptogium cyanescens, magnifed 40X, with lobule-shaped isidia.

An isidium (plural: isidia) is a minute, cylindrical or branched outgrowth found on the thallus of certain lichen species. Isidia primarily function as a means of vegetative propagation, allowing the lichen to spread and colonize new substrates. It is a diaspore–a reproductive propagule comprising both fungal and algal components. An isidium mirrors the anatomical structure of the parent thallus from which it arises, in that it is surrounded by a cortex. It is this latter feature that contrasts it with soredia, a similar reproductive structure that lacks a cortex. In some cases, the term isidium is used rather broadly, to include all propagules with a cortex, even ones that are only externally similar, such as dactyls or pseudoisidia.

Isidia have a diverse morphology, ranging in shape from spherical and cylindrical to club-shaped, with sizes typically between 0.01 to 0.03 mm in diameter and 0.5 to 3.0 mm in height. Their appearance can vary, being smooth to knobby or shiny to matte, with some lichens featuring hollow, inflated-looking isidia. Less often, isidia are compressed and spathulate, squamose (scaly), or even shield-like ("peltate"). Differentiating between isidia and other lichen outgrowths can be challenging, as they can resemble juvenile isidia or other thalline structures. While some isidia detach easily or transform with age, others remain permanently attached to the thallus, likely increasing its surface area. Isidia that have broken off can be carried by wind, water, or animals (such as getting stuck on bird feet or entangled in animal fur). When these land in a conducive environment, they can give rise to new thalli. While isidia are still connected to a thallus, they enhance its surface area, thereby aiding in gas exchange and photosynthesis.

About 25–30% of foliose (leafy) or fruticose (bushy) lichens form isidia.[1] Features of isidia are reflected in the species epithets of many lichens, both indirectly, and explicitly.

Morphology[edit]

Closeup of the pustulate isidia that are characteristic of Flavoparmelia baltimorensis; scale bar = 1 mm

The visual characteristics of isidia can show a high degree of variability. These tiny protrusions have been observed to manifest as spherical (globose), cylindrical, scale-like (squamulose, coralloid,[2] club-shaped (claviform), disc-shaped (disciform), cup-shaped (cupuliform),[3] and wart-shaped verruciform.[4] Their size generally ranges from 0.01 to 0.03 mm in diameter and 0.5 to 3.0 mm in height.[5] Their colour, particularly at the apex, often differ from non-isidiate sections. They vary from simple to branched. The surface can be smooth to knobby, and shiny to matt.[4] While most isidia are solid in composition, a few lichens feature isidia that are hollow and have an inflated appearance. The formation of soralia and isidia can display some overlap. Soralia can grow at the tips of isidia, and isidia can develop within a soralium. In the latter case, the loose network of the soralium undergoes a compact outgrowth that then develops a cortex secondarily.[2]

The isidia of the foliose lichen Pseudocyphellaria horridula stand out distinctly. They emerge prominently from the lobes in a vertical manner and can grow quite large – reaching lengths of up to 10 mm. These isidia, possibly the largest observed in lichens, impart a distinctive fish bone pattern to the lobes.[6]

Parmelina pastillifera produces a unique type of isidia. This foliose species has a greyish colour with a central, subsenescent part that is covered by button-shaped protuberances. These protuberances are relatively large, brown to black, and have a flat or slightly concave, warty upper surface. They are easily broken off, leaving a crater-shaped depression in the mother thallus' surface. This feature makes it possible to differentiate P. pastillifera from P. tiliacea, another greyish foliose species that has much smaller, globose to cylindrical, black isidia on the central, older parts of the thallus, which are morphologically very similar.[7]

the gelatinous lichen Collema subflaccidum has prominent globular isidia.

It can be challenging to differentiate between isidia and other outgrowths and appendages of the thallus since certain lichens exhibit similar-looking warts and papillae on their surface, which resemble juvenile isidia. Old isidia might show only slight differences from foliose outgrowths, known as lobuli. Additionally, there are gradual transitions to fibrils and small lateral branches. In certain lichens, old isidia can transform into other thalline structures without breaking off. Only a few lichens have isidia that are actively dispersed by the degeneration of their cortical base. The isidia of many crustose lichens, such as the Pertusariaceae, can be easily detached. Others can only be freed by the death of the thallus. In many gelatinous and foliose lichens, the isidia stay attached to the thallus permanently. In such lichens, they likely serve the purpose of increasing the surface area.[2] Consoredia are an aggregation or cluster of incompletely separated soredia; soralia with an abundance of consoredia may be superficially mistaken for aggregations of isidia.[8] In some cases, however, the isidia of normally isidiate species may break down to a mass of soredia and consoredia. Some species, such as Pertusaria coccodes, may rarely produce both isidia and soredia.[9] The hybrid term "sorediate isidia" has been applied to isidia that bear or erupt into soredia, usually at the tips. Similarly, "isidiate (or isidioid) soredia" has been used to describe soredia that resemble isidia in that they are often darkened and solid looking, but lack a cortex and often arise from distinct soralia. Du Rietz used the term "isidiate soredia" to describe the delicate and readily fragmented isidia of Xanthoparmelia loxodes and X. verruculifera,[10] but as pointed out by Theodore Esslinger, the terminology was inappropriate, as no soredia are actually formed.[11] The ability of soredial granules to grow out like isidia on the parent thallus has been noted to be quite common in the Physciaceae.[12]

Formation and development[edit]

Isidia proliferate on the surface of this Pseudevernia furfuracea
Isidia are abundant on the lobe margins of this Peltigera praetextata

The formation and development of isidia are diverse, influenced by various tissue initiations and environmental conditions. In many lichens, the genesis of isidia starts with the intrusion of medullary hyphae or those from the algal layer between the cortical hyphae. These hyphae transport algal cells upwards, leading to a protuberance that initially lacks a differentiated cortex, forming secondarily as the process matures. This mode of development is notably prevalent in pseudocyphellae, where the outgrowing hyphae aren't constrained by a cortex.[2]

Expanding our knowledge further, experimental studies on Pseudevernia furfuracea have unveiled the intricacies of isidia formation. It is found to involve high cellular turnover in both symbiotic partners, marked by an upsurge in asexual spore production of algae and the proliferation of surrounding medullary hyphae. Such isidia are metabolically vibrant, as evidenced by their heightened photosynthesis and robust dark respiration rates.[13]

Transitioning to the specifics of Peltigera praetextata, in this lichen, isidia emerge solely at sites of cortical injury. Such development can be experimentally instigated by making physical incisions on the thallus. However, the terminology used to label these foliose outgrowths remains contentious. Some researchers propose the term "lobuli" as a more fitting descriptor, emphasizing their regenerative role.[2]

Venturing into another formation strategy, some isidia stem from a protuberance of the thalline cortex. The tissue from the algal layer advances into this protrusion. Remarkably, in certain scenarios, isidia originate solely from cortical hyphae that ensnare free-living algae. These algae become encapsulated by the outgrowing hyphae of the cortex. Subsequent connections to the internal layers of the thallus are secondary in nature. Such a formation process is evocative of cephalodia, highlighting the adaptability inherent in lichen tissues.[2]

Lastly, delving into the world of gelatinous lichens, isidia's birth is set in motion by the active division of algal cells at the thallus's periphery. For those without a cortex, this triggers a minuscule protuberance that is soon invaded by hyphae. Meanwhile, in corticate ones, a spike in algal proliferation spurs the cortical cells into heightened division. This orchestrated growth ensures that the resultant thalline protuberance is uniformly coated with a cortical layer from the get-go.[2]

Regeneration[edit]

While many isidia might not primarily contribute to vegetative propagation, they play a pivotal role in regeneration. On aging thalli, isidia can sprout into new lobes. This phenomenon was exemplified in a study observing Parmotrema tinctorum isidia. When these isidia were kept in nylon bags on Cryptomeria japonica trunks, they began to form protuberances within six months, eventually maturing into lobules with rhizines and lichen compounds by the year's end.[14] Additionally, the isidia of the cyanobacterial species Peltigera praetextata, when sown onto soil, formed small juvenile thalli in 4–5 months, with larger thalli appearing after eight months.[15]

A commonly held perspective suggests that post-dissemination, isidia do not transform directly into a full thallus. They are believed to first deconstruct into a loosely tied association of fungal and algal cells, which subsequently forms a new thallus.[2] However, evidence from certain species contradicts this idea. The isidia of Phyllophiale pastillifera, for instance, are adapted for rapid thallus development shortly after detachment.[7] Similarly, the isidia of Lobaria pulmonaria also appear to germinate directly into young thalli without the need for a preliminary dedifferentiation phase.[16]

In terms of reproduction, while the relatively hefty weight of isidia makes them less conducive to dispersion than soralia, they have the advantage of resilience in adverse conditions, thanks to their protective cortex. Though larger lichen fragments can result in swifter regeneration, their mass complicates dispersal.[2]

For the foliose lichen Parmelia saxatilis, mature isidia evolve into fresh thalline scales on the thallus surface. These emergent scales rejuvenate growth by overlapping older thalli in a tile-like fashion, utilizing the tissue from older, deceased lobes as a water reservoir.[2]

Transplantation studies[edit]

Lichen transplantation is a restoration technique employed to introduce or reintroduce lichens to areas where they've been depleted or to new environments where they can be beneficial. This method aims to enhance ecosystem recovery and may serve various purposes, from environmental remediation to conservation efforts. Isidia, given their role in lichen propagation, have become valuable assets in such transplantation studies.

For instance, isidia from the foliose lichen Xanthoparmelia tinctina were used in an attempt to rehabilitate an abandoned asbestos mine. The goal was to introduce lichens to cover and stabilize the surface, reducing the exposed asbestos surface area and thereby mitigating the dispersal of hazardous airborne fibers. Unfortunately, achieving robust colonization was a challenge, with running water and debris posing significant hurdles.[17]

From a conservation perspective, Parmotrema crinitum isidia were cleverly combined with surgical gauze fibers to facilitate effective substrate attachment during transplantation. On the other hand, Sticta sylvatica required a considerable 24-month period post-transplantation to generate small lobules from its isidia.[18]

Function and ecological role[edit]

  • Role of isidia in lichen reproduction.
  • Advantages of having isidia for lichen colonization and distribution.
  • Interaction with surrounding environment and other organisms.

Vegetative propagules like isidia can probably germinate in a broader range of habitats than ascospores or conidia, since they do not require a resynthesis of the symbiosis.[19]

Leptotrema and Graphina rely on isidia to facilitate the aeration of their thallus. The apex of the isidia in these species contains hyphal tissue that is made porous and loosened, providing a transition point to a respiratory pore. When the isidia detach, they leave behind a respiratory pore that resembles a crater. This pore can be sealed by a secondary cortex that forms during subsequent development.[2]

Types and variations[edit]

Isidia that occur on Usnea, such as these on the branch S. ceratina, are sometimes referred to as isidomorphs.

Isidia present in various forms and structures, each providing unique insights into lichen biology and taxonomy. In his 1992 monograph of the bark-dwelling, sorediate and isidiate crustose lichens of Norway, Tonsberg broadly defined isidia as "all globose to elongate, usually projecting, corticate diaspores with a basal point of attachment", and further defined several types.[4] He delineated these as follows:

The first group comprised isidia that were initiated beneath the upper cortex and seemed to burst through the thallus cortex. Such isidia, continuously developed, pushed older ones outward with younger isidia forming beneath. They lacked a primary cortex, with cortex differentiation being secondary. This resulted in homoiomerous isidia, meaning their interiors contained both hyphae and algae without a medullar central core. The mature isidia were surrounded by ecorticate tissue in rounded patches, often located at tuberculae tips. This was observed in species like Pertusaria flavida, P. coccodes, and P. coronata.[4]

Pseudevernia consocians has claviform (club-shaped) isidia.[3]

Some isidia connected continuously with the surrounding thallus cortex and seemed to originate from it. This was seen in species like Pertusaria corallina, P. oculata, and P. dactylina. These isidia were heteromerous, meaning they had a white medullary core surrounded by an algal layer. This formation mode differs anatomically from the homoiomerous isidia of the foliose species Lasallia pustulata and Parmelia tiliacea, which are also derived from the thalline cortex.[4]

A few areolate species displayed isidiiform areolae, which are more or less spherical to cyclindrical, and easily detached or broken. Examples include Caloplaca herbidella and Placynthiella icmalea. The latter's isidia-like areolae were termed "blastidia", as they resembled consoredia but remained attached at the base to the substrate. The entire areola of Placynthiella icmalea was identified as a blastidium.[4]

An isidiomorph is a structure that resembles an isidium, but is formed as an outgrowth of the medulla rather than the cortex. It is associated with soralia of species in the genus Usnea.[20]

Schizida and thlasidia[edit]

The terms schizidia and Thlasidia were introduced by Josef Poelt to categorize specific diaspores and isidia-like structures in lichens. Introduced in 1965, schizidia describe flattened diaspores formed when the lichen thallus's upper layers split, often resembling scale-like, rounded segments, exemplified in Baeomyces rufus. Conversely, Thlasidia, presented in 1986, pertains to the cylindrical, easily detached structures of Gyalideopsis anastomosans, which mirror isidia and function likely as diaspores.[4]

In Heiomasia sipmanii, Aptroot et al. (2009) labeled disc-shaped isidioid structures as schizidia, deviating from the schizidia definition given by Frisch & Kalb (2006) for the genus Stegobolus within the Graphidaceae family. These structures in H. sipmanii and H. seaveyorum are more like thallus outgrowths than true isidia, the latter defined by an upper cortex and photobiont layer.[21] Terms such as "pseudisidia" introduced by Aptroot and colleagues[22] and "pseudoisidia" can lead to confusion. The former refers to isidioid structures without a cortex, while the latter signifies isidia-like formations devoid of photobiont cells.[21] Considering the ambiguities, Nelsen et al. suggest referring to these Heiomasia structures simply as isidia, regardless of their lack of cortex — an attribute stemming from the thallus being ecorticate, not an intrinsic feature of isidia.[23]

The debate over these terminologies remains, especially when discussing ecorticate isidia-like outgrowths in lichens like Heiomasia seaveyorum. "Pseudisidia" and "pseudoisidia" have been interchangeably employed by different authors across taxonomic groups, furthering the confusion.[24]

Another variation, an isidioid soredium, manifests as a secondary corticate protuberance in soralia-like clusters. Notably, polyisidia, which are clustered isidia formed on thalline outgrowths, are exclusive to the genus Pyxine.[21] Thlasidia, on the other hand, bear a resemblance to pseudoisidia at their terminal ends but house photobiont cells in soredia-like patches at their bases. Originating from the thlasidium, they are solely reported in the crustose, epiphytic lichen Gyalideopsis anastomosans.[21]

Another unique structure is the Phyllophiale-type isidium, sometimes referred to as scutelliform isidia.[25] This disc-shaped propagule, first linked with the foliicolous lichen fungus Phyllophiale (now part of Porina), has a small Phycopeltis alga thallus encircled by fungal hyphae. This structure grows via its hyphal fringe and algal filaments, forming a fungal network over adjoining algal thalli. This eventually leads to the production of similar disc-like propagules elevated from the main thallus.[25]

Taxonomic value[edit]

Erik Acharius was the first to define a genus (Isidium) based on the presence of this trait. William Nylander defined many species of Parmelia based on the presence or absence of isidia. Du Rietz's 1924 classification of isidia ascribed basic taxonomic value to these formations, and he was the first to develop the idea of species pairs–morphologically and chemically identical but differing in the presence or absence of vegetative diaspores.[12] In certain genera, isidia attributes serve as vital taxonomic markers. For instance, Leptogium lichens can be differentiated based on their isidium development and the morphology of mature isidia. Two primary isidium developmental types are recognized: hirsutum-type and saturninum-type. Furthermore, mature isidia within certain Leptogium species offer clues for distinguishing superficially similar species due to unique characteristics they exhibit.[26]

However, the value of isidial features in taxonomy has been contested. Poelt noted that the formation of isidia in the genus Collema, which was used by Gunnar Degelius to delineate varieties and forms, was "little stabilized morphologically or systematically".[12] In another example, a study by Kalb and Hafellner's described Porina isidiata as the isidiate counterpart of specimens resembling P. atlantica, an otherwise identical species. Observations in various regions indicated that P. isidiata might simply be a variant of P. atlantica. In the genus Porina, distinguishing between genuine isidia and abnormal photobiont outgrowths has proven challenging. McCarthy (1993) and Harris (1995) posited that some of these structures might result from aggressive photobiont growth. Concurrently, certain species, notably P. ocoteae, displayed isidioid outgrowths in stressed conditions. These and similar observations suggest that the presence of "isidia" should not be a defining taxonomic criterion in all cases, as differences in the structure of these propagules may be influenced by environmental factors.[27]

Distribution and habitat[edit]

  • General habitats where lichens with isidia are commonly found.
  • e.g. this, from the 2023 paper: “It seems that, in groups of perithecioid lichens, isidia are much rarer than in apothecioid lichens (e.g. Diederich and Ertz (2020); Orange and Chhetri (2022))”
  • Factors influencing the distribution of isidia-bearing lichens.

Lichens that produce vegetative propagules commonly occur without apothecia, or, if they do occur, they are often sparse or immature.[28] It has been estimated that 25–30% of foliose or fruticose lichens form isidia.[1]

Eponyms[edit]

Niebla isidiascens is one of many lichens named for the taxonomic uniqueness of their isidia.

In lichens, specific features related to their isidia are often captured in their species epithet. For instance, the epithet of Porina coralloidea alludes to its isidial features,[29] while Aspicilia stalagmitica earned its name due to distinct isidia-like outgrowths.[30]

Many authors incorporate the term "isidia" (or its variations) directly in species epithets, emphasizing its taxonomic significance. Here, a range of these eponyms are listed, with each accompanied by its respective author citation and year of publication:

Astrothelium isidiatum, characterized as a new species in 2023, is the only species of the large family Trypetheliaceae that is known to produce isidia.[31]

Furthermore, some species names combine "isidia" with descriptive terms to portray specific characteristics:

Acanthotrema alboisidiatum is named for the white isidia that contrast with its light olive-green thallus.

In some cases, species are named for both their isidial attributes and geographical distribution, exemplified by epithets such as N. amerisidiata, N. australisidiata, N. brasilisidiata, and N. siamisidiata.[43]

History[edit]

Erik Acharius was the first to recognise the taxonomic value of the coral-like outgrowths he saw on the thalli of some lichens.[44] In 1794, he circumscribed the genus Isidium with the following diagnosis: "branchlets produced on the surface, or coralloid, simple and branched".[45] The genus included densely isidioid species such as Isidium corallinum (now Lepra corallina) and I. westringii (now Pertusaria pseudocorallina). Genus Isidium has since been synonymised with Pertusaria.[46]

The development of isidia was described by German botanist Friedrich Rosendahl in several species of Parmelia.[47] In one of them, P. papulosa, which has a cortical layer one cell thick, the isidium begins as a small swelling or wart on the upper surface of the thallus. At that stage the cells of the cortex have already lost their normal arrangement and show irregular division. They divide still further, as gonidia and hyphae push their way up. The full-grown isidia in this species are cylindrical or clavate, simple or branched. They are peculiar in that they bear laterally here and there minute rhizoids, a development not recorded in any other isidia. The inner tissue accords with that of the normal thallus and there is a clearly marked cortex, gonidial zone and pith. A somewhat analogous development takes place in the isidia of Parmelia proboscidea; in that lichen they are mostly prolonged into a dark-coloured cilium.[44]

In Parmelia scortea the cortex is several cells thick, and the outermost rows are com- pressed and dead in the older parts of the thallus; but here also the first appearance of the isidium is in the form of a minute wart. The lower layers (4 to 6) of living cortical cells divide actively; the gonidia also share in the new growth, and the protuberance thus formed pushes off the outer dead cortex and emerges as an isidium. They are always rather stouter in form than those of P. papulosa and may be simple or branched. The gonidia in this case do not form a definite zone, but are scattered through the pith of the isidium.[44]

Here also should be included the coralloid branching isidia that adorn the upper surface and margins of the thallus of Umbilicaria pustulata. They begin as small tufts of somewhat cylindrical bodies, but they sometimes broaden out to almost leafy expansions with crisp edges. Most frequently they are situated on the bulging pustules where intercalary growth is active. Owing to their continued development on these areas, the tissue becomes slack, and the centre of the isidial tuft may fall out, leaving a hole in the thallus which becomes still more open by the tension of thalline expansion. New isidia sprout from the edges of the woundand the process may again be repeated. It has been asserted that these structures are only formed on injured parts of the thallus something like gall formations but Bitter has proved that the wound is first occasioned by the isidial growth weakening the thallus.[44] Du Rietz wrote about isidia shape about 100 years ago, categorizing them according to shape: globose, cylindrical, claviform, or coralloid.[10] The term was first used in the sense it is used now by Georg Meyer (botanist) [de] in 1825, and adopted by Elias Fries in 1831.[48]

The term isidium is derived from the New Latin term isidium, which itself comes from the Ancient Greek isis, Classical Greek: Ἶσις, meaning "coral", with the diminutive suffix idium.[3] The adjectival form of isidia is isidiate.[3] Also, the term isidiiferous refers to a lichen thallus bearing isidia.[49]

To track the microscopic reproductive propagules of lichens, a low-cost device was designed to trap them from the air. Molecular techniques were concurrently developed to enable species-specific identification of the captured propagules, with the capability of detecting a single asexual propagule, such as an isidium. This combined approach of using mechanized propagule traps with DNA diagnostics represents a pioneering effort to study lichen dispersal.[50]

References[edit]

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Cited literature[edit]

Tønsberg, T. (1992). The Sorediate and Isidiate, Corticolous, Crustose Lichens in Norway. Sommerfeltia. Vol. 14. p. 129. ISBN 82-7420-015-2.

Category:Lichenology Category:Fungal morphology and anatomy

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