Mediterranean Cladoniaceae A
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Mediterranean Cladoniaceae A. R. Burgaz, T. Ahti & R. Pino-Bodas Spanish Lichen Society (SEL) Madrid 2020 Mediterranean Cladoniaceae Ana Rosa Burgaz, Teuvo Ahti & Raquel Pino-Bodas Spanish Lichen Society (SEL) Madrid 2020 Mediterranean Cladoniaceae Ana Rosa Burgaz (Complutense University, Madrid) Teuvo Ahti (Finnish Museum of Natural History, Helsinki) Raquel Pino-Bodas (Royal Botanic Gardens, Kew) Photographer: Pablo Galán-Cela (Polytechnic University, Madrid) Cladoniaceae Supported by research project CGL2013-41839-P. Ministry of Science, Innovation and Universities, Spain. With the collaboration of Prof Soili Stenroos (Finnish Museum of Natural History, Helsinki), Dr Edit Farkas and Dr László Lőkös (Hungarian Academy of Sciences and Hungarian Natural History Museum, Budapest), Dr Mohammad Sohrabi (Iranian Research Organization for Science and Technology, Tehran), Dr Ayhan Şenkardeşler (University of Ege, Izmir, Turkey) and under the auspices of the Spanish Lichen Society (SEL) www.ucm.es/info/seliquen © Sociedad Española de Liquenología (SEL) Depósito Legal: B-18067-2020 ISBN: 978-84-09-21610-9 Impresión: erasOnze Artes Gráficas Cladoniaceae Introduction The Old World Mediterranean Region lies in Southern Central Italy have a temperate climate in the lower levels, Europe, Southeast Asia and Northern Africa, surround- while above the treeline the climate is alpine (for more ing the Mediterranean Sea basin (Fig. 1). Although it information see Nimis 2016). Most of Morocco, Algeria, represents only 1.6% of the land area of the planet, 10% Tunisia and Libya present desert climates. of all known vascular plants grow in this region, 50% of The Mediterranean lichens have been more in- which being endemic (Cowling et al. 1996). The domi- tensely studied in some countries for which a complete nant vegetation in this region is of sclerophyllous type, lichen flora or checklist exist. These countries are France characterized by perennial, often small-sized leaves, (Ozenda & Clauzade 1970; Roux et al. 2014, 2017), Italy usually covered with waxes. These woodlands have been (Nimis 1993, 2016; Nimis & Martellos 2017; Nimis et al. managed for centuries to obtain pastures, resulting in 2018), Greece (Abbott 2009; Linda 2019), Spain and agroforestry ecosystems characterized by scattered trees Portugal (Llimona & Hladun 2001). An important con- (21-40% of canopy) and an annual grass layer, with a sa- tribution to the Western European lichen flora was pub- vannah-like landscape. The succesional stages of these lished by Clauzade & Roux (1985). The origin of many forests are dominated by shrubs of Fagaceae, Lamiaceae and of these studies was an international project to prepare Cistaceae or Pinus halepensis formations (San-Miguel-Ayanz a checklist of the Mediterranean lichens (Nimis 1996). et al. 2016), with different designations, depending on the The cited works were forerunners of the study of the country. Species of genera with a paleotropical origin, family Cladoniaceae in this region. Subsequently, mono- such as Myrtus, Phillyrea and Pistacea, are also frequent. graphic studies of the family, or of the genus Cladonia, A transition to main nemoral European broadleaved as well as new records, have been published for the forests occurs in the mountains and is constituted by Iberian Peninsula (Burgaz & Ahti 2009; Pino-Bodas et al. warm-temperate deciduous trees (Quercus spp., Fraxinus 2013b, 2014), Greece (Sipman & Ahti 2011), Bosnia- ornus, Ostrya carpinifolia or Acer spp.) which occupy sub- Herzegovina and Croatia (Burgaz & Pino-Bodas 2012), mediterranean locations (Box & Fujiwara 2015). The re- Italy (Gheza et al. 2018), Turkey (Kocakoya et al. 2018), gion is considered one of the world biodiversity hotspots etc. However, still now the knowledge on the Cladoniaceae (Blondel & Aronson 1999) and the study of its flora has in the Mediterranean basin is much less complete than in appealed many researchers (Maya et al. 2017). The paleo- Central and Northern Europe, where the distribution of geologic and climatic events that have succeeded each most of the species is well known (Ahti & Stenroos 2013). other through time, along with the great orographi- This is the reason why we decided to carry out the study cal heterogeneity and the soil variety are responsible of the family Cladoniaceae for the countries surrounding for the region’s high floristic wealth. The dominant the Mediterranean Sea, in order to compile a catalogue as climate of the countries in this region is the so-called complete as possible of the species present in the region, Mediterranean climate, characterized by the aridity of adding distribution maps of the species and data about the summer season, often a prolonged absence of rain- their chemical variation. fall. Summer temperatures tend to exceed 35 ºC, while The area under active study embraced the follow- winters are usually mild, although the continental areas ing countries or regions: Portugal, Spain, Andorra, Italy, may be subject to severe frosts. The western areas are Southern France, Malta, Croatia, Bosnia-Herzegovina, more humid through the influence of the Atlantic Ocean. Montenegro, Albania, Greece, Cyprus, Turkey, Morocco, Nevertheless, a narrow strip with Atlantic climate is lo- Algeria, Tunisia and Libya. All the authors have per- cated in the Northern Iberian Peninsula and much of formed extensive field work in Spain and Portugal, but France. The Alps in Northern Italy and the Apennines in in order to carry out the present survey field work new 6 mediterranean cladoniaceae Fig. 1. Countries surrounding the Mediterranean basin. Modify from https://d-maps.com/. Alb: Albania; B&H: Bosnia-Hercegovina; Kos: Kosovo; Mol: Moldova; Mon: Montenegro; N.Ma: North Macedonia; Slov: Slovenia ; TRNC: Turkish Republic North Cyprus. collections were made by the author Burgaz in some of the has examined material in numerous other herbaria in less surveyed countries: Albania, Greece, Malta, Croatia, course of many years, especially Paris (PC), London Montenegro and Cyprus. In addition, new collections were (BM), Stockholm (S), Uppsala (UPS), Vienna (W, WU), made in Southern France, Southern Italy and Sardinia. Berlin (B), Geneva (G) and Copenhagen (C). The species The author Ahti has independently collected Cladoniaceae distribution maps have been drawn main using the ma- in Sardinia, NE Italy, Greece and Turkey (there with Pino- terial studied by Burgaz along with some bibliographic Bodas). From the herbaria or literature we also added the quotations; the sources are distinguished on the maps by presence of each species in San Marino, Egypt, Lebanon, means of different symbols, ▲( ) for the specimens stud- Syria and Israel (including all Palestine). ied and (●) for literature reports. This study is meant as Besides many Spanish and Portuguese herbaria, a compendium of the knowledge on Cladoniaceae in the several other herbaria have been visited to achieve a countries that border the Mediterranean for a world more complete sampling: Florence (FI), Helsinki (H), monograph. It provides new records for several countries Budapest (BP), Tbilisi (TBI). In addition, the author Ahti and extends the distribution of a number of species. Cladoniaceae The family Cladoniaceae comprises about 500 species dis- at the apex of the podetia, more rarely they are sessile on the tributed in 18 genera (Stenroos et al. 2019), only three squamules or on a short stipe. The colour of the hymenium of them present in Europe, Cladonia, Pilophorus and varies from dark, brown to almost black to light brown, but Pycnothelia. Most of the species (ca. 475) belong to the ge- red colour hymenia also appear. The asci show an apical nus Cladonia (Ahti 2000; Wijayawardene et al. 2018). The amyloid thickening (I+), with a clearer central channel, and first stage of development begins with the formation of a are surrounded by a strongly amyloid tube of Porpidia-type. prothallus, constituted by the hyphae from the germina- They contain 8 ascospores, which are simple, rarely with tion of an ascospore. Almost immediately the protothallus 1-3 septa, hyaline, from fusiform to ovoid. The paraphyses contacts the alga, starting the lichenization and develop- are simple and septate. The characters associated with the ing small squamules of dorsiventral structure which con- apothecia lack taxonomic value in most of the genera. stitute the primary thallus. Most of the taxa have a mixed Anatomically, the thallus have a compact ectal layer thallus, consisting of two parts, one basal, parallel to the called cortex, lacking in many species, made up of hyphae substrate, called primary thallus and other erect, the sec- with short cells, with thickened walls and vertical orien- ondary thallus. The secondary thallus consists of vertical tation. The surface of the cortex is variable and has tax- structures, fruticulose, hollow, exceptionally solid, called onomic value. Sometimes an epinecral layer outside the podetia (when it is constituted by generative tissue) or cortex exists. Under the cortex the algal layer appears, pseudopodetia (when it is constituted by vegetative tis- which can be continuous or discontinued by photobiont sue). The taxonomy of the family is mostly based on the glomerules. Immediately below the algal layer the medul- morphology of these structures, which can range from la is located, formed by lax hyphae. In many species, the simple to very complex branching patterns. boundary between the algal layer and the medulla is not The tips of the podetia may be linear,