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- 🧬 Over 50% of lichen species thought to be widespread contain cryptic species; this means biodiversity estimates are twice what we thought.
- 🌍 Molecular phylogenetics shows regional endemism and their hidden past in lichens that look the same.
- 🧪 DNA barcoding, especially of the ITS region, is now the main way for identifying lichen-forming fungi species.
- ⚖️ Misidentifying cryptic species can hurt bioindicator accuracy and conservation strategies.
- 🔍 Integrative taxonomy using molecular, chemical, ecological, and morphological data gives strong species classification.
The Hidden Diversity in Lichen-Forming Fungi
Lichens, the crusty, colorful life forms seen on rocks, bark, and soil, are one of nature’s most fascinating experiments in symbiosis. Composed of a fungal partner and a photosynthetic ally (usually green algae or cyanobacteria), lichens are close ecological units. However, new research shows that these partnerships hide a lot of unseen diversity—what scientists call “cryptic species.” Thanks to progress in molecular phylogenetics, researchers are looking past what they look like, into their DNA. This is showing a complex family tree that greatly changes what we know about biodiversity.

What Are Cryptic Species?
Cryptic species are groups of organisms that are genetically distinct but look almost the same. Traditional taxonomy, which mainly looked at what things looked like, often misses the small differences that separate these species. This oversight has long affected the study of various organisms, especially fungi and lichen-forming fungi. Here, form can look the same even when genetic groups are not related.
When studying fungi, identifying species just by how they look often makes things too simple. A lichen species commonly found on temperate tree bark may look identical to one in a tropical mountain range. Yet DNA sequencing tells another story—these look-alike organisms may be different family lines that split apart millions of years ago.
This misidentification is not just a small academic point—it affects real-world applications. This ranges from species conservation to the trustworthiness of bioindicator data used to monitor environmental health. Understanding and correctly identifying cryptic species are thus very important steps in ecological science, managing biodiversity, and even public policy.

Why Lichens Are a Perfect Place for Cryptic Diversity
Lichen-forming fungi are prone to having many cryptic species due to a mix of biological traits and how they interact with their surroundings, which makes it hard to tell them apart visually.
Symbiotic Complexity
Lichens are composite organisms resulting from a symbiosis between fungi (the mycobiont) and photosynthetic partners (the photobiont), creating a new form unlike either partner alone. Multiple genetically distinct fungi can produce lichens that look almost the same when paired with similar photobionts. This forms what is known as “photomorphs”—species that differ genetically but look identical because they have the same partners.
These partnerships can be different in various groups or places, even if they look the same from the outside. In addition, some fungi can partner with multiple photobionts in different places. This makes their looks even more varied, confusing older ways of classifying them.
Environmental Plasticity
Lichens are very adaptable, changing how they look and their color depending on things in their environment like moisture, light, and air quality. A lichen species in a humid coastal forest might look fuller and greener than the same species growing in an arid plateau, where it looks shrunken and bleached. This plasticity makes it hard to compare how they look, and makes it seem like different species are actually the same one.
Convergent Morphology
Convergent evolution is when unrelated species get similar traits because they face similar environmental conditions. In lichenology, this is common—different species from very different family lines sometimes grow in the same way, or have the same color or shape, because they adapt to similar conditions. As a result, just looking at how they appear can greatly undercount the real number of different species.
Together, these factors make lichen-forming fungi a perfect place for cryptic species—not only are they complicated biologically, but their past makes these differences hard to see.

Molecular Phylogenetics: The DNA Revolution
What Is Molecular Phylogenetics?
Molecular phylogenetics is the study of evolutionary relationships by analyzing DNA sequences. Instead of identifying organisms based on what they look like (phenotype), it looks at how their DNA is related to other organisms'. For lichen-forming fungi, this has been very telling.
Key Tools and Methods
- ITS sequencing: The Internal Transcribed Spacer (ITS) region of ribosomal DNA works like a “barcode” for species identification among fungi. It’s the main standard and most commonly used molecular marker.
- Multilocus sequence analysis (MLSA): This means sequencing many genes or parts of the genome. It gives clearer answers for family lines that are hard to figure out.
- Next-generation sequencing (NGS): Allows for fast, large-scale sequencing of entire genomes or environmental DNA samples. It is good for finding unknown cryptic species even in mixed samples.
- Phylogenetic trees: Scientists map the evolutionary relationship between DNA sequences in a diagram known as a tree. This shows splits and differences, which points to many species.
According to research by Lumbsch and Leavitt (2010), these tools have changed how lichenized fungi are classified, moving from just looking at how they appear to using strong genetic proof. This DNA-first approach has already found many cryptic family lines we would not have seen otherwise.

Case Studies: When DNA Split Familiar Faces
Molecular studies have changed how we define familiar lichen species across the globe.
The Parmelia saxatilis Complex
Once considered a single species that could have many forms, Parmelia saxatilis is now known to be a group of genetically different family lines found across different continents. Though visually similar, these family lines show big genetic differences and also live in different places and like different conditions. The result: One species became many—each with its own story of how it changed over time.
Lecanora polytropa Reclassified
Another good example is Lecanora polytropa, a crustose lichen once seen as a common species found all over the world. Molecular phylogenetics showed this was not true. Genetic analysis found multiple cryptic groups, each living in its own type of environment—alpine habitats, volcanic rock outcrops, and arctic tundra—and perhaps changed over time to have special chemical interactions with the surfaces they grow on.
These discoveries don’t just highlight species misidentification. They make us look at older ecological studies in a new way, changing what we thought about lichen ecology, geographic distribution, and how they adapted.

Naming Cryptic Species: A Taxonomic Tightrope
The process of naming a newly discovered species is full of tough questions and practical problems, especially when it comes to cryptic species.
Model Paradoxes: MSC vs. PSC vs. BSC
- Morphological Species Concept (MSC): Classifies species based on observable traits. This method fails for cryptic species, which don’t show clear physical differences.
- Phylogenetic Species Concept (PSC): Defines species based on unique genetic lineages. It is great at finding cryptic species, but it might separate species too much based on small genetic differences.
- Biological Species Concept (BSC): Looks at whether species can reproduce with each other. This is often hard to apply in fungi where testing mating compatibility is not practical.
With molecular phylogenetics more and more used, many taxonomists think we should change species definitions to rely on genetic differences, especially when also using ecological or chemical information.
Consequences of Proper Taxonomy
Accurate naming is not just academic—it helps guide environmental rules, conservation funding, and international databases like GBIF and GenBank. Incorrectly lumped species can receive the wrong protection or be completely missed for protection.
Making taxonomy clear also helps businesses, for example, when looking for new uses for lichens used in traditional medicine, cosmetics, or natural dyes. Knowing what species you’re working with improves both how well things work and how lasting they are.

Undiscovered Riches: How Many Cryptic Species Remain?
Estimates vary, but the hidden diversity among lichen-forming fungi is probably huge.
According to Crespo and Pérez-Ortega (2009), more than 50% of lichen species found all over the world, once they were looked at again with genetic tools, were actually groups of many species. This means that thousands of species we thought were single could actually be tens of thousands if studied correctly. If we do this for all fungi, we could find twice or even three times as many fungal species as we know now.
This new count changes conservation targets, ecological surveys, and our overall understanding of life on Earth.

Lines of Evidence: Identification Beyond Genetics
While DNA analysis is powerful, no single method guarantees exact species definitions. Today’s best practice is integrative taxonomy—using many fields of study to define species.
Core Evidence Streams:
- Morphology: Modern microscopes and analytic software can spot tiny body differences that match genetic family lines.
- Molecular Data: DNA remains the main way to measure—especially ITS and other conserved regions.
- Chemistry (secondary metabolites): Lichen species often produce special chemicals that we can find using methods like thin-layer chromatography or HPLC.
- Ecology/Niche profiling: Some cryptic species live in certain heights, on specific surfaces, or in particular weather conditions, showing they are different species.
When all these proofs come together, cryptic species are seen as real and are clearly placed in the classification system.

Phylogeography: Mapping Cryptic Trends Across Areas
Phylogeography connects where organisms live to how they changed over time, showing how cryptic species spread out in different natural areas.
For example:
- Glacial refugia: Regions that remained ice-free during the last Ice Age often were safe places for family lines that later grew. This led to small genetic differences between groups that moved back into different world regions.
- Mountain ranges and valleys: Physical blocks to genetic mixing cause them to split, which makes cryptic species that are used to separate, small weather conditions.
- Oceanic islands and archipelagos: Isolated by water, lichens change on their own even if they look the same as ones on the mainlands.
These processes add to species only being found in one region, and show even more biodiversity.

Impacts on Conservation and Bioindicators
Mistaken species identities can seriously hurt conservation efforts. A “widespread” lichen found to be many rare cryptic species that only live in small areas needs local conservation plans, not general ones.
Similarly, many lichens are bioindicators—organisms used to measure air quality, radiation, or contamination. If what’s thought to be a single indicator for all places is actually a dozen different species with different levels of what they can handle, the conclusions drawn from such studies could be wrong or not true.
Clarifying cryptic diversity thus makes conservation biology and ecological checks more exact, leading to better funding, focused action, and new priorities.

Why Mushroom Growers and Curious Citizens Should Care
Even outside academic circles, the discovery of cryptic species matters in practical ways.
Improved Cultivation Targeting
Understanding evolutionary relationships can help grow the kinds of fungi we want or fungi that are tough, helping indoor growers, permaculturists, and medicinal researchers find the best fungi to use.
Commercial Applications
Cryptic family lines may produce special natural chemicals used in antibiotics, cosmetics, or plant probiotics that can handle tough conditions. Identifying them opens new ways for research and development.
Amplifying Scientific Participation
There’s more and more push for citizen science tools that can spot cryptic species. With smartphone lenses, field notebooks, or eventually, handheld DNA tools, non-specialists could help add to worldwide fungus databases.

Future Tech: DNA for the People
The tools that found cryptic species are quickly becoming cheaper and more accessible.
- Portable DNA Sequencers: Devices like the Oxford Nanopore MinION mean we can now do DNA analysis outside the lab.
- AI-Powered Identification: Using machine learning with huge classification data sets lets computers identify fungi from pictures, making it more exact.
- Community Databases: Platforms like iNaturalist and Mushroom Observer keep adding more lichen sightings—AI and molecular uploads are the next big step.
With tools everyone can use and more awareness, everyday nature lovers can become the next big helpers in classifying species.
Peeling Back Nature’s Layers
Cryptic species in lichen-forming fungi are an amazing reminder that nature rarely shows everything it has. Through molecular phylogenetics, we learn that even the most familiar organisms may have many unseen identities. These discoveries go beyond just academic study—changing conservation rules, updating how we classify species, and making our understanding of life’s complexity deeper.
In a world changing faster than ever, finding hidden diversity is not just a scientific job, but a shared duty. Whether you’re looking through a microscope, hiking a lichen-covered trail, or growing a mushroom kit at home—there’s always a mystery waiting beneath the surface.
References
Bickford, D., Lohman, D. J., Sodhi, N. S., Ng, P. K. L., Meier, R., Winker, K., Ingram, K. K., & Das, I. (2007). Cryptic species as a window on diversity and conservation. Trends in Ecology & Evolution, 22(3), 148–155. https://doi.org/10.1016/j.tree.2006.11.004
Crespo, A., & Pérez-Ortega, S. (2009). Cryptic species and species pairs in lichens: A discussion on the relationship between molecular phylogenies and morphological characters. Mycologia Balcanica, 6(1-2), 121–127.
Lumbsch, H. T., & Leavitt, S. D. (2010). Goodbye morphology? A paradigm shift in the delimitation of species in lichenized fungi. Fungal Diversity, 38, 45–58.



