Fossil Usnea and similar fruticose lichens from Palaeogene amber

Fruticose lichens of the genus Usnea Dill. ex Adans. (Parmeliaceae), generally known as beard lichens, are among the most iconic epiphytic lichens in modern forest ecosystems. Many of the c. 350 currently recognized species are widely distributed and have been used as bioindicators in air pollution studies. Here we demonstrate that usneoid lichens were present in the Palaeogene amber forests of Europe. Based on general morphology and annular cortical fragmentation, one fossil from Baltic amber can be assigned to the extant genus Usnea. The unique type of cortical cracking indirectly demonstrates the presence of a central cord that keeps the branch intact even when its cortex is split into vertebrae-like segments. This evolutionary innovation has remained unchanged since the Palaeogene, contributing to the considerable ecological flexibility that allows Usnea species to flourish in a wide variety of ecosystems and climate regimes. The fossil sets the minimum age for Usnea to 34 million years (late Eocene). While the other similar fossils from Baltic and Bitterfeld ambers cannot be definitely assigned to the same genus, they underline the diversity of pendant lichens in Palaeogene amber forests.Peer reviewe

Symbionts and changing environment: Lichen diversity and photobiont associations in tropical mountain ecosystems

Epiphytes comprise a significant component of biodiversity and biomass in tropical forests. They are ecologically important in intercepting and retaining moisture, providing habitat and food for invertebrates, and contributing fixed nitrogen into the ecosystem. Lichens are mutualistic symbioses between lichen-forming fungi (mycobionts) and algae and/or cyanobacteria (photobionts). Most lichen mycobionts are specific in their photobiont choice and the local availability of compatible photobionts may limit their ability to disperse into new habitats. The aims of this study are to 1) provide the first account of lichen symbiont diversity in tropical mountains, with focus on changes along topographic gradients, and 2) elucidate the effects of human induced environmental change to lichen symbiotic organisms, including the effects of expansion of agricultural and other disturbed ecosystems, and changing climate. The results will be a significant contribution to understanding tropical biodiversity since so far very few studies deploying modern molecular biological methods have included lichens from East Africa. Lichens, bryophytes, and free-living cyanobacteria and green algae, along the natural environmental gradient of the southern slope of Kilimanjaro including all main ecosystem types. The sampling is focused on study plots established by the KiLi project . The collected specimens will be studied microscopically, with chemical analyses, and molecular biology methods. So far we have sampled several plots within the natural savanna, maize fields, grassland, and Chagga homegardens (3–5 sampled plots each ecosystem type). The specimens have been studied microscopically. The preliminary results show, that clear differences exist in lichen biota between different plot types: lichen abundance seems to depend especially on presence/absence of woody plants, lichen species on the climate, and lichen diversity on substrate variability. In all studied plots lichens mainly occur epiphytically on shrubs and trees.Non peer reviewe

A botanical view of the ‘Baltic amber forest’: new evidence from seed plants, lichens and fungi

Baltic amber forms the largest amber deposit on earth and it is particularly well-known for the plethora of arthropod inclusions. The floristic composition, habitat types and climate of its Eocene source area, however, are still controversial. The differing suggestions range from early Eocene tropical to late Eocene temperate environments, and from lowland to montane forests. We screened a large number of inclusions from historic collections and from recently discovered amber pieces and found many inclusions of seed plants, lichens and microfungi that provide important insights into habitat structure and climate.Peer reviewe

Taitaia, a novel lichenicolous fungus in tropical montane forests in Kenya (East Africa)

During lichenological explorations of tropical montane forests in Kenya, a remarkable new lichenicolous fungus was repeatedly found growing on thalli of the epiphytic tripartite cyanolichen Crocodia cf. clathrata. Molecular phylogenetic analyses placed the fungus within Gomphillaceae (Ostropales, Lecanoromycetes), a family mainly of lichen-symbiotic species in the tropics. The anatomical features (unitunicate, non-amyloid asci and simple, septate paraphyses) as well as the hemiangiocarpic ascoma development confirm its taxonomic affinity. DNA sequence data showed the closest relationship was with Gyalidea fritzei, followed by Corticifraga peltigerae. A monotypic genus, Taitaia, is introduced to incorporate a single species, T. aurea. The new fungus is characterized by aggregated ascomata with yellow margins and salmon red discs developing from a single base.Peer reviewe

Evolution of the tRNALeu (UAA) intron and congruence of genetic markers in lichen-symbiotic Nostoc

The group I intron interrupting the tRNALeu UAA gene (trnL) is present in most cyanobacterial genomes as well as in the plastids of many eukaryotic algae and all green plants. In lichen symbiotic Nostoc, the P6b stem-loop of trnL intron always involves one of two different repeat motifs, either Class I or Class II, both with unresolved evolutionary histories. Here we attempt to resolve the complex evolution of the two different trnL P6b region types. Our analysis indicates that the Class II repeat motif most likely appeared first and that independent and unidirectional shifts to the Class I motif have since taken place repeatedly. In addition, we compare our results with those obtained with other genetic markers and find strong evidence of recombination in the 16S rRNA gene, a marker widely used in phylogenetic studies on Bacteria. The congruence of the different genetic markers is successfully evaluated with the recently published software Saguaro, which has not previously been utilized in comparable studies.Peer reviewe

Complex interaction networks among cyanolichens of a tropical biodiversity hotspot

Interactions within lichen communities include, in addition to close mutualistic associations between the main partners of specific lichen symbioses, also more elusive relationships between members of a wider symbiotic community. Here, we analyze association patterns of cyanolichen symbionts in the tropical montane forests of Taita Hills, southern Kenya, which is part of the Eastern Afromontane biodiversity hotspot. The cyanolichen specimens analyzed represent 74 mycobiont taxa within the order Peltigerales (Ascomycota), associating with 115 different variants of the photobionts genus Nostoc (Cyanobacteria). Our analysis demonstrates wide sharing of photobionts and reveals the presence of several photobiont-mediated lichen guilds. Over half of all mycobionts share photobionts with other fungal species, often from different genera or even families, while some others are strict specialists and exclusively associate with a single photobiont variant. The most extensive symbiont network involves 24 different fungal species from five genera associating with 38 Nostoc photobionts. The Nostoc photobionts belong to two main groups, the Nephroma-type Nostoc and the Collema/Peltigera-type Nostoc, and nearly all mycobionts associate only with variants of one group. Among the mycobionts, species that produce cephalodia and those without symbiotic propagules tend to be most promiscuous in photobiont choice. The extent of photobiont sharing and the structure of interaction networks differ dramatically between the two major photobiont-mediated guilds, being both more prevalent and nested among Nephroma guild fungi and more compartmentalized among Peltigera guild fungi. This presumably reflects differences in the ecological characteristics and/or requirements of the two main groups of photobionts. The same two groups of Nostoc have previously been identified from many lichens in various lichen-rich ecosystems in different parts of the world, indicating that photobiont sharing between fungal species is an integral part of lichen ecology globally. In many cases, symbiotically dispersing lichens can facilitate the dispersal of sexually reproducing species, promoting establishment and adaptation into new and marginal habitats and thus driving evolutionary diversification.Peer reviewe

An annotated checklist of the bryophytes of Taita Hills region, Kenya

Based on previous literature and our own collections, we list 288 bryophyte species (145 liverworts, 143 mosses) from the Taita Hills region (including Mt. Kasigau and Maktau Hill) in SE Kenya. New records for Kenya include the liverworts Archilejeunea elobulata Steph., Bazzania nitida (F. Weber) Grolle, Cololejeunea grossepapillosa (Horik.) N. Kitag., Diplasiolejeunea kraussiana (Lindenb.) Steph., D. villaumei Steph., Lejeunea amaniensis E.W. Jones, L. cyathearum E.W. Jones, Lopholejeunea laciniata E.W. Jones, Metzgeria crassipilis (Lindb.) A. Evans, M. nudifrons Steph., Plagiochila boryana (F. Weber) Nees, and P. moenkemeyeri Steph., and the mosses Leucophanes hildebrandtii Müll. Hal. and Neckeromnion lepineanum (Mont.) S. Olsson, Enroth, Huttunen & D. Quandt. A further 22 liverworts and 13 mosses previously known from other parts of Kenya are reported for the first time from the Taita Hills region.Peer reviewe

Lichen preservation in amber : morphology, ultrastructure, chemofossils, and taphonomic alteration

The fossil record of lichens is scarce and many putative fossil lichens do not show an actual physiological relationship between mycobionts and photobionts or a typical habit, and are therefore disputed. Amber has preserved a huge variety of organisms in microscopic fidelity, and so the study of amber fossils is promising for elucidating the fossil history of lichens. However, so far it has not been tested as to how amber inclusions of lichens are preserved regarding their internal characters, ultrastructure, and chemofossils. Here, we apply light microscopy, scanning electron microscopy (SEM), energy dispersive X-ray spectroscopy (EDX), and Raman spectroscopy to an amber-preserved Eocene lichen in order to gain information about the preservation of the fossil. The lichen thallus displays lifelike tissue preservation including the upper and lower cortex, medulla, photobiont layer, apothecia, and soredia. SEM analysis revealed globular photobiont cells in contact with the fungal hyphae, as well as impressions of possible former crystals of lichen compounds. EDX analysis permitted the differentiation between halite and pyrite crystals inside the lichen which were likely formed during the later diagenesis of the amber piece. Raman spectroscopy revealed the preservation of organic compounds and a difference between the composition of the cortex and the medulla of the fossil.Peer reviewe

Sensitivity of tropical pendant bryophytes : results from a translocation experiment along an elevation gradient

We studied growth of three epiphytic bryophyte species in Kenya to determine their sensitivity to environmental changes. Bryophytes collected from a cool and moist upper montane forest were used to prepare 180 pendant transplants. Sixty transplants were placed in their natural habitat while 120 were transferred to two warmer and drier sites in a lower montane forest. After one year, all the transplants recovered from the lower mon-tane forest were transferred back to the upper montane forest. In the third year, half of the remaining transplants were left in their location, and half transferred to an eucalyptus plantation forest, both in the upper montane zone. After each year subsamples were taken from each group for growth measurements. The epiphytic bryophytes demonstrated considerable resilience during the experiment. They clearly suffered from the warm and dry conditions of the lower montane forest, but quickly recovered and then exhibited growth rates comparable to those of the controls. In the third year there was no statistical difference in the growth of transplants in their natural location and in the eucalyptus plantation. This indicates that the lack of suitable substrate is the primary factor limiting bryophyte biomass in plantation forests rather than unsuitable microclimate per se.Peer reviewe