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The origins and evolution of island floras as exemdified by the AGgean archipelago
... In addition to the previous example, the work of Greuter (1972Greuter ( , 1979Greuter ( , 1995-a systematic botanist-on island taxa indicated that island taxa, and insects predominate in these assemblages, exhibit some particularities. First, the earlier the habitation of the island by humans, as it happens in the Aegean and the Mediterranean in general, the lower the extinction rates unlike the islands in the Pacific where the human occupation varies inversely with extinction rates (Pimm et al. 1995). ...
... Birds adapt rapidly to human occupation and sparrows, starlings and doves are good examples of such an anthropophily while plants -and insects as well -are adapted to human settlements only through pre-adaptive characteristics and evolution of new traits. However, the evolution in the Aegean islands is almost at a standstill (Greuter 1979) and in this sense the island floras are of relict type except from some 'cryptoendemics', which form very sparse populations unlikely to be exploited by insects. The transgression of sea level in the interglacials lowered the temperature on the mountains by 6-7 o C and sea surface by 4 o C, which indicates that certain plant taxa will become extinct together with their arthropod faunas. ...
... Contrary to the findings of Ribera and Vogler (2004) for diving beetles, islands have never been evolutionary theaters and no plant originated in there since the Messinian salinity crisis. This entire region, Cardaegean region according to Greuter (1979) remained insular throughout Pleistocene sea transgressions. All these facts assure that extinction rates cannot be extrapolated in a geographical or a taxonomical sense, while in every case there is a sort of fragmentation of the previous geographical range of insect taxa. ...
The question 'how many species inhabit the earth?' is not just the result of the inherent property of human mind for continuous intellectual search. It is the urgent need to know the biodiversity of our planet, to predict its fate and increase our understanding of various biological species, their mutual dependencies and their abiotic requirements. Insects constitute the dominant taxonomic group of the earth. Many authorities summing both the taxonomically described and undescribed species, estimate the total to be from 1.8 to 50 millions (May 1988, 1990; Stork 1993). In an attempt to assess the ratio of the described to undescribed living taxa Hammond (1990, 1992) showed that insects are the richest taxonomic group in both categories. Even the described species surpass all the other groups of organisms in all five kingdoms. In this respect it is expected that among the organisms that are going to become extinct in the next few decades insects will constitute a large proportion (Stork 1993). Pimm and Raven (2000) estimate that among living species one-tenth may become extinct in the next fifty years. This is expected because of extensive habitat loss, which in a wider sense includes habitat fragmentation and inbreeding suppression among the remaining populations. Despite the fact that insects, especially the strict specialists, are among the first organisms liable to become extinct our knowledge on their taxonomy, distribution and ecological role is very limited. What we need is time and money, for the documentation of insect extinctions (Dunn 2005). In this context the monitoring of insect extinctions is different from that of mammals and birds. Dunn (2005) states that the IUCN Red List (2002) records 129 bird species extinctions within the last 600 years, which constitute 1.3% of all bird species. By analogy we may expect through the world some 44,000 of the estimated total of 3.4 million insect species to become extinct. However, only 70 insect species extinctions have been documented so far. This great discrepancy is largely the result of our limited study of insects. If we accept the arguments of other authors in biodiversity calculus and the estimation of insect species this discrepancy becomes even larger. The 'guestimates' of world insect species derived by Stork (1993) is 5.3 million species. Gaston (1992) estimates range between 6.88 and 8.75 million species. Hodkinson and Casson (1991) extensively collected tropical Hemiptera species sampled with all known devices from Sulawesi, Borneo estimated the number of insect species of the world to be 3.0 - 4.0 millions. On the same line of arguments by accepting the working figure of 10,000 beetle species in the collection area and taking species accumulation curves the estimates rise to 5.0 - 6.0 million species. Other much higher numbers, though not so convincing are provided by Erwin (1992) who estimated the world insect fauna as 30.0 million, while on the basis of distribution models relating species abundance and body size of various taxa inferred global estimates of insect fauna between 10.0 - 50.0 million (May 1988, 1990). Surprisingly, the number of studies predicting the richness of the world insect fauna is not balanced with studies predicting the number of insect species to be affected by modern extinctions either on a global or a regional scale. One explanation lies probably in the fact that science is dominated by western schools of thought and associated cultures in which insects and arthropods in general are primarily considered as pests that should be controlled (Kim 1993a, b). An immediate consequence is the exclusion of insects from almost all conservation plans, except possibly those for butterflies and some dragonflies. In the current extinction event, which is induced by humans, the need to separate background extinctions from exceptional extinctions is urgent since it leads to the quantification of local extinctions and the organization of a global network of databases and experts. It is well known to perceptive field entomologists that many insect populations are disappearing from their area of collection and that other species may invade the area establish new populations or even a network of metapopulations depending on the fragmentation status of the habitat. Nevertheless, it is not known if local extinctions are part of a global mass extinction or they are merely above-background events. Paleontologists used to face the problem by interpreting the scarcity or disappearance of insect families as extinctions in excavation layers corresponding to geologic epochs (Labandeira 2005), a fact that reflects the efforts and the cost rather than scientific reality. In this chapter I have attempted to present the extent of current insect extinctions and relate them to possible taxonomic bias. Also, causes of extinctions and similarities with previously recorded mass extinctions are discussed. Current theories about colonization of an area by insects and the applicability of these to current insect extinctions and their detection is examined. The necessity for extensive taxonomic studies is obvious and the ways that this can be achieved and function in the estimation of species loss are presented.
... The "native nonendemics," sometimes termed "natives," are a widely spread taxonomic group occurring both on the Aegean islands and on the mainland across the Mediterranean (2,673 taxa), ~40% of which reached the Aegean islands as a result of human action in prehistoric times (Greuter, 1971(Greuter, , 1979Kougioumoutzis et al., 2020, mod-ern introductions were excluded). The "multiple continental Greek endemics" (GE) are species exclusively occurring both on the Greek mainland and the Aegean islands (689 taxa). ...
... However, we did not investigate the issue further as it goes beyond the scope of this study. For angiosperms, the lack of influence of biogeographic affinity can partially be attributed to the fact that a large portion (~40%) of the present Aegean flora has reached the Aegean islands due to human action in prehistoric or early historic times (Greuter, 1971(Greuter, , 1979Kougioumoutzis et al., 2020). For reptiles, the lack of effect of biogeographic affinity could result from similar levels of diversity in the various source communities "seeding" the different islands east and west of the Aegean Trench (Foufopoulos et al., 2011). ...
Recent research in island biogeography has highlighted the important role of late Quaternary sea‐level fluctuations in shaping biogeographic patterns in insular systems but focused on oceanic systems. Through this study, we aim investigate how late Quaternary sea‐level fluctuations shaped species richness patterns in continental‐shelf island systems. Focusing on the Aegean archipelago, we first compiled maps of the area's geography using published data, under three sea‐level stands: (a) current; (b) median sea‐level over the last nine glacial–interglacial cycles (MSL); and (c) Last Glacial Maximum (LGM). We gathered taxon–island occurrences for multiple chorotypes of angiosperms, butterflies, centipedes, and reptiles. We investigated the impact of present‐day and past geographic settings on chorological groups by analyzing island species–area relationships (ISARs) and using generalized linear mixed models (GLMMs) selection based on multiple metrics of goodness of fit. Our results confirm that the Aegean's geography has changed dramatically since the LGM, whereas the MSL only modestly differs from the present configuration. Apart for centipedes, paleogeographic changes affected both native and endemic species diversity through altering connections between land‐bridge islands and the mainland. On land‐bridge islands, we detected over‐representation of native species and under‐representation of endemics. Unlike oceanic islands, sea‐level‐driven increase of isolation and area contraction did not strongly shape patterns of species richness. Furthermore, the LGM configurations rather than the MSL configuration shaped patterns of endemic species richness. This suggests that even short episodes of increased connectivity with continental populations are sufficient to counteract the genetic differentiation of insular populations. On the other hand, the over‐representation of native nonendemic species on land‐bridge islands reflected MSL rather than LGM mainland connections. Our study shows that in terms of processes affecting species richness patterns, continental archipelagos differ fundamentally from oceanic systems because episodic connections with the mainland have profound effects on the biota of land‐bridge islands.
... The impressive morphological polymorphism among these species led to the recognition of many intraspecific taxa. Eddie & Ingrouille (1999) presented a detailed morphological study, especially on the trilocular C. rupestris complex and the quinquelocular C. andrewsii and C. lyrata complexes ( Fig. 1A and 1H, respectively), suggesting that the high diversity in leaf and flower morphology may represent a mosaic of rapidly evolving populations that are correlated with the geological and climatic history of the Aegean, especially the eustatic sealevel changes during the three million years of the Pliocene and the climatic fluctuations of the Pleistocene (Greuter, 1979). Our findings provide evidence for the young age of these species, supporting the hypotheses of Eddie & Ingrouille (1999), and these radiations were likely favored by small-scale vicariant events created by the complex paleogeographical history of the Aegean. ...
... In the case of Campanula chasmophytic species, rainstorms and wind may play some role in range expansion ), but long-distance dispersal does not seem a plausible process to explain the fragmented distribution of Campanula in the Aegean. Greuter (1979) documented the flora of the island of Psara where the only population of C. merxmuelleri is found, growing on a wall of a monastery (Baliousis, 2016), and noted that 45% of the island flora has an anthropogenic origin. Therefore, recent introductions cannot be excluded. ...
Campanula sect. Quinqueloculares (Campanulaceae), consisting of ca. 39 mostly chasmophytic species, is one of the most
morphologically variable groups in Campanula and includes numerous endemics occurring mostly in Greece and/or Turkey. In this
molecular study, we aim to test the monophyly of C. sect. Quinqueloculares and provide divergence time estimates to generate
hypotheses into the historical processes responsible for the diversification and current distribution patterns of this group. Individual
and combined data matrices consisting of plastid (NADHS-2, rpoC1-1, rpoC2-1, rpoC2-2, rpoC2-3, trnT-trnL) and nuclear (2017561,
ITS, PPR11, PPR70) markers were constructed for 121 taxa. Results indicate that C. sect. Quinqueloculares, as traditionally circumscribed,
is polyphyletic. Species are largely clustered into two well-supported clades, except for three taxa excluded from these
groups. Additionally, a few taxa belonging to other sections are confidently nested within the two Quinqueloculares clades. The first
clade (Greek clade) includes one isophylloid species nested with 25 Greek endemics belonging to C. sect. Quinqueloculares. The second
clade (Southeastern Aegean-Turkish clade) comprises 20 C. sect. Quinqueloculares taxa and 3 species of C. sect. Rupestres, all
distributed in the southeastern Aegean and Turkey. Divergence time estimates suggest that these clades originated in the Late Miocene.
Temporal and geographic patterns are consistent with a vicariant scenario driven by geological events during the Miocene, such
as the formation of the Mid-Aegean trench and the Messinian salinity crisis.
Keywords Aegean archipelago; Campanula sect. Quinqueloculares; Campanulaceae
... This is not surprising, since a noteworthy proportion of plant diversity and Mediterranean endemic taxa are located on the > 10 000 islands and islets in the basin (Médail, 2017). To trace the origin and evolutionary history of island species, it is crucial to understand the background of island formation and its putative past geological connection with mainland areas (Greuter, 1979;Fernández-Palacios et al., 2016). ...
... In the Mediterranean Basin two main island types can be found: continental islands, which were separated from the continental masses by rising sea level and/or tectonic processes and, less commonly, oceanic islands, which formed de novo by volcanic activity (Greuter, 1979). Continental islands can harbour species coming from the same sources as those on oceanic islands (i.e. from a continental source or from other neighbouring islands), but also species that evolved from ancient continental lineages that were separated by tectonic plate movements. ...
To what extent Pleistocene sea-level fluctuations have affected the genetic diversity of species is one of the current topics in biogeographical research. Carduncellus dianius is a Mediterranean narrow endemic species, restricted to < 20 populations distributed along coastal areas in Alicante (mainland eastern Iberian Peninsula) and on the island of Ibiza (Balearic Islands). To get insights into its evolutionary history and its genetic diversity and structure, we combined the analysis of molecular markers (three plastid DNA regions and AFLP) with ecological niche modelling. Results from dated phylogeographical analyses revealed that this species might have originated in the continental region during the early Pleistocene. The colonization of Ibiza could have occurred by a single long-distance dispersal event, with a subsequent back-colonization from the island to the same continental area of origin. These results corroborate the role of islands as sources for mainland colonization (biodiversity reservoirs) and as refugia during glacial periods. Notably, we detected that populations located on stable landmasses (i.e. not affected by sea rising during interglacial cycles) harboured significantly higher genetic diversity than those that were periodically submerged during the periods of marine transgressions. Our results point out sea-level fluctuations as a factor to be considered in phylogeographical studies focused on species distributed along coastlines.
... Every island is unique in terms of geographical and topographical features, e.g., position, size, altitude, geodiversity, origin, geohistorical processes and many more (Tzanoudakis & Panitsa, 1995). Enduring human impact is combined with the geographical features in larger islands to shape the insular flora (Greuter, 1975a, b;Whittaker & Fernández-Palacios, 2007); this is especially true for the Aegean islands, since, nearly 45% of the present Aegean flora has reached the Aegean Islands owing to human action in prehistoric or early historic times (Greuter, 1979). ...
... The most prominent of these factors are discussed below. Sfenthourakis and Triantis (2017) underlined that the understanding of the interplay between human presence, establishment of exotic species and extinction of indigenous biotas, is of critical importance.Human influence is long-lasting in the Aegean as in the whole Mediterranean area and human factor has significantly shaped the Aegean flora since synanthropic floristic element reaches 30%-50% of some island floras and it is the most effective vector for long-distance dispersal (Greuter, 1975b(Greuter, , 1979(Greuter, , 1995(Greuter, , 2001Cellinese et al., 2009;Triantis & Mylonas, 2009, Kallimanis et al. 2010, Trigas, Panitsa, & Tsiftsis, 2013. Kougioumoutzis and Tiniakou (2014) suggested that the human factor has played a major role in shaping the pattern observed in the endemic species present in the Cyclades. ...
The Aegean archipelago is one of the largest archipelagos in the world and has long fascinated biogeographers due to its high environmental heterogeneity, complex palaeogeography, high diversity and endemism. In this study, prominence has been given to the plant diversity and biogeography of the Aegean area. After describing the phytogeographical aspects in the Aegean archipelago from the first division in phytogeographical areas, the current widely used subdivisions to the recent aspects of the Aegean phytogeographical classification, a synoptic analysis of the Aegean plant diversity is presented focusing on the Aegean endemics, the range-restricted plant taxa, the single island endemics, the protected plant taxa and the most Critically Endangered ones. Plant diversification and speciation in the Aegean and factors driving them, as revealed by molecular studies, are discussed briefly. The ruderal, alien and invasive plant taxa richness has also been mentioned. Emphasis is also given to small islands and islets plant species diversity. Factors affecting plant species richness in the Aegean, such as the long-lasting human presence, climate, area, elevation, habitat diversity, isolation, geological substrate and structure are discussed on the basis of different biogeographical studies concerning the Aegean area.
... Every island is unique in terms of geographical and topographical features, e.g., position, size, altitude, geodiversity, origin, geohistorical processes and many more (Tzanoudakis & Panitsa, 1995). Enduring human impact is combined with the geographical features in larger islands to shape the insular flora (Greuter, 1975a, b;Whittaker & Fernández-Palacios, 2007); this is especially true for the Aegean islands, since, nearly 45% of the present Aegean flora has reached the Aegean Islands owing to human action in prehistoric or early historic times (Greuter, 1979). ...
... The most prominent of these factors are discussed below. Sfenthourakis and Triantis (2017) underlined that the understanding of the interplay between human presence, establishment of exotic species and extinction of indigenous biotas, is of critical importance.Human influence is long-lasting in the Aegean as in the whole Mediterranean area and human factor has significantly shaped the Aegean flora since synanthropic floristic element reaches 30%-50% of some island floras and it is the most effective vector for long-distance dispersal (Greuter, 1975b(Greuter, , 1979(Greuter, , 1995(Greuter, , 2001Cellinese et al., 2009;Triantis & Mylonas, 2009, Kallimanis et al. 2010, Trigas, Panitsa, & Tsiftsis, 2013. Kougioumoutzis and Tiniakou (2014) suggested that the human factor has played a major role in shaping the pattern observed in the endemic species present in the Cyclades. ...
The Aegean archipelago is one of the largest archipelagos in the world and has long fascinated biogeographers due to its high environmental heterogeneity, complex palaeogeography, high diversity and endemism. In this study, prominence has been given to the plant diversity and biogeography of the Aegean area. After describing the phytogeographical aspects in the Aegean archipelago from the first division in phytogeographical areas, the current widely used subdivisions to the recent aspects of the Aegean phytogeographical classification, a synoptic analysis of the Aegean plant diversity is presented focusing on the Aegean endemics, the range-restricted plant taxa, the single island endemics, the protected plant taxa and the most Critically Endangered ones. Plant diversification and speciation in the Aegean and factors driving them, as revealed by molecular studies, are discussed briefly. The ruderal, alien and invasive plant taxa richness has also been mentioned. Emphasis is also given to small islands and islets plant species diversity. Factors affecting plant species richness in the Aegean, such as the long-lasting human presence, climate, area, elevation, habitat diversity, isolation, geological substrate and structure are discussed on the basis of different biogeographical studies concerning the Aegean area.
... It is noteworthy that the earliest archaeological evidence for Neolithic economies in SE Europe dates to c. 7000 cal BC, with the founding of a fully-fledged farming community at Knossos on the island of Crete [58]. It has been estimated that approximately one third of the wild flora of Crete has been introduced by man [49,59,60]. Moreover, human pressure and overgrazing has probably altered the present distribution patterns of many plant species [56,61]. ...
... The low altitude peak of species richness on Crete may have also been further supported by human transferred species, since human activities are mainly confined to lowland areas. Plant species already introduced in prehistoric or early historic times may be perfectly integrated into the native plant communities and it may be extremely difficult, or even impossible, to distinguish them from the truly native ones [60]. ...
Elevational ranges of the differentiated island-mainland endemics (DIME) distributed in Crete and the Peloponnese and the resulted IMAR for each species.
(DOCX)
... According to my results, it does indeed seem to play an important role, but perhaps not in the same way as for oceanic islands, which are highly invasible ecosystems par excellence. The mechanisms used to explain oceanic island invasibility often include niche availability due to taxonomic disharmony characteristic of highly isolated communities (Greuter 1979;Mueller-Dombois & Loope 1990;Givnish 1998;Lonsdale 1999). But the small, offshore islands studied here, even though they do have an insularity syndrome as concerns richness and diversity (Médail & Vidal 1998), are not comparable to an oceanic island in terms of taxonomic disharmony (see also Greuter 1979). ...
... The mechanisms used to explain oceanic island invasibility often include niche availability due to taxonomic disharmony characteristic of highly isolated communities (Greuter 1979;Mueller-Dombois & Loope 1990;Givnish 1998;Lonsdale 1999). But the small, offshore islands studied here, even though they do have an insularity syndrome as concerns richness and diversity (Médail & Vidal 1998), are not comparable to an oceanic island in terms of taxonomic disharmony (see also Greuter 1979). A key characteristic of these little islands that might better explain their invasibility is the fact that their fauna is dominated by certain species that occur in extremely high densities, and constrained to a small surface area. ...
Chapter 1: Plant invasions: introduction to a global phenomenon
Chapter 2: Carpobrotus spp as a biological model for the study of plant invasions in the Mediterranean Basin
Chapter 3: Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: I. Genetic diversity and introgression
Chapter 4: Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: II. Reproductive strategies
Chapter 5: Unexpected insularity effects in invasive plant mating systems: the case of Carpobrotus (Aizoaceae) taxa in the Mediterranean Basin
Chapter 6: The evolutionary potential of invasive Carpobrotus (Aizoaceae) taxa: Are gene flow and hybrid vigor connected?
Chapter 7: Invasion-associated hybridization and change: a karyological and morphological comparison of native and Mediterranean Carpobrotus (Aizoaceae) taxa
Chapter 8: The impacts of Carpobrotus spp. (Aizoaceae) on native plant communities in southeast France
Chapter 9: The pollinators of native and introduced plants: Does co-evolution matter?
Chapter 10: Invasional meltdown potential: facilitation between introduced plants and mammals on French Mediterranean islands
Chapter 11: Discussion
... The numerically small but diverse campanulaceous taxa of North America probably contain many relicts from pre-glacial times and represent several relatively independent groups derived from the main rapunculoid radiation in Eurasia (Shetler, 1979 ). An early radiation of the Rapunculus group in the Northern Hemisphere would explain the distinctiveness of subgroups (e.g., Phyteuma, Petromarula, and related genera) that are associated with the European Alpine orogenic events and fluctuating Mediterranean sea levels during the Tertiary period (Eddie, 1984; Favarger, 1972; Greuter, 1979). It would also explain the presence of endemic genera such as Githopsis in California and the other rather heterogeneous taxa in North America, e.g., Heterocodon and diverse Campanula annuals in California (see Morin, 1980 ), China, and southern Asia (e.g., Homocodon D. Y. Hong and Peracarpa J. D. Hooker & T. Thoms.). ...
... clade remain unclear (see also Hilliard & Burtt, 1973). Symphyandra A. DC. is now generally considered to be artificial (Greuter et al., 1984; Oganesian, 1995 ), and this analysis supports that conclusion . However, the four sections of the genus recognized by Fedorov (1957) are all quite distinct, and we suggest that the species formerly included in this genus should be re-examined and not necessarily included in Campanula without substantial evidence. ...
Ninety-three taxa comprising thirty-two genera (plus four outgroups from Lobeliaceae) were used to estimate a phylogeny of the Campanulaceae based on ITS sequences of nuclear ribosomal DNA. From 2629 most parsimonious trees, a strict consensus tree with bootstrap values was constructed, in addition to a phylogram showing branch lengths. The topologies of these two trees are discussed in relation to the pollen and capsule morphology within the family, in addition to chromosome number and geographical distribution. The results show that there is a major dichotomy between the colpate/colporate pollen alliance (platycodonoid taxa) and the porate pollen alliance (wahlenbergioid and campanuloid taxa). Both these major alliances are monophyletic. Within the porate alliance there are two major clades, the wahlenbergioids and the campanuloids. The campanuloid clade is further subdivided into two major clades representing the Rapunculus and the Campanula s. str. groups of taxa, plus three smaller clades that are considered as ''transitional'' taxa. It is argued that the family originated in a fragmenting West Gondwanaland and that tectonic processes are responsible for the major dichotomy in the family. The colpate/colporate platycodonoids subsequently remained relatively relictual in Asia, whereas the porate taxa spread over much of the Northern and Southern Hemispheres. The campanuloid lineage spread over the Northern Hemisphere from a major evolutionary center in the Mediterranean region and is represented in North America only by the Rapunculus group. The wahlenbergioid lineage is widely dispersed across the southern continents and oceanic islands but has a major secondary center of diversification in southern Africa. The use of ITS provides insights for future investigations and a phylogenetic framework that can be tested with other data sets. Its limitations for phylogeny reconstruction are briefly discussed. More extensive taxon sampling and additional data sets are required to refine these results and for a new classification of the Campanulaceae to be proposed.
... Good examples for the explanation of recent distribution patterns of plants and animals in the eastern Mediterranean area were and are the subject of various studies. Some of the earlier works by CREUTZBURG (1963) or GREUTER (1970GREUTER ( , 1979, as well as more recent publications by FATTORINI (2002FATTORINI ( , 2006, CHATZIMANOLIS et al. (2003) or POULAKAKIS et al. (2015), should be mentioned here. The sometimes-significant human intervention through development or grazing, which has led to severe impoverishment not only on the smaller islands (BERGMEIR & DIMOPOULOS 2003), is worthy of consideration. ...
Summary
Hirth, M. & H.F. Paulus (2024): Pollination biology, morphology, taxonomy of
the small flowered species of the Ophrys holosericea aggregate in the East
Aegean.- J. Eur. Orch. 56 (2-4): 219-266.
For many years, we have been visiting several of the East Aegean islands to
study the orchid flora there. For this work, we paid particular attention to the
different populations of the small-flowered Ophrys holosericea group, which
presented themselves differently on each of the islands visited. As smallflowered,
we group together plants of the Ophrys holosericea aggregate with lip
length less than 1 cm. To understand these better and, above all, to be able to
establish their taxonomic-systematic status, we visited the larger islands of
Samos and Kos in particular, as well as the smaller islands of Phournoi, Tilos
and Lipsi. To be able to establish the populations concerned as a biospecies, we
studied the pollination biology in particular. The following results can be
presented.
1. On the island of Samos, we discovered the presence of two different “small
flowered holosericeas”. One in the center, north of the city of Mytilene, with
a substantial population in an extensively used vineyard. The other mainly in
the southwest of the island, in the Limnionas-Agia Kyriaki area. Both forms
differ in flower morphology and have different pollinators. The vineyard
population has already been published in advance in the new field guide by
KREUTZ (2024) under the name Ophrys ampelaki M.Hirth, Paulus, Kreutz. Its
pollinator is the small long-horned bee Eucera digitata. The other form is
Autorenexemplar - copyright by AHO Baden-Württemberg
220 Journal Europäischer Orchideen 56 (2-4): 2024.
described as O. kerkissoula M.Hirth & Paulus in this article. This species also
occurs on the neighbouring island of Phournoi. Its pollinator is Eucera
laxiscopa.
2. From the small island of Lipsoi. a small flowered holosericea is reported but
not finaly discribed, as the data base is insufficient and the pollinator is
unknown.
3. Ophrys tili from the island of Tilos was re-examined and compared with the
very similar-looking plants from Kos which had already been described as
O. ellinicaea Kreutz & B. Tenschert (2011). Since both forms are hardly to
distinguish and both have the same pollinator, Eucera cineraria, we consider
them to be conspecific.
4. In a phylogeographic context, the current distributions of the species are
discussed with regard to data presented by KOUGIOUMOUTZIS et al. (2017) on
pre- and postglacial land changes in the eastern Aegean.
... Gute Beispiele für die Erklärung rezenter Verbreitungsmuster von Pflanzen und Tieren im östlichen Mittelmeerraum waren und sind Gegenstand verschiedener Untersuchungen. Zu nennen sind hier einige frühe Arbeiten von CREUTZBURG (1963) oder GREUTER (1970GREUTER ( , 1979, sowie neuere Veröffentlichungen von FATTORINI (2002FATTORINI ( , 2006, CHATZIMANOLIS et al. (2003) oder POULAKAKIS et al. (2015. Bedenkenswert sind die zum Teil erheblichen menschlichen Eingriffe durch Bebauungen oder Beweidung, die zu empfindlichen Verarmungen nicht nur auf den kleineren Inseln geführt haben (BERGMEIR & DIMOPOULOS 2003). ...
changes in the eastern Aegean.
Zusammenfassung
Hirth, M. & H.F. Paulus (2024): Bestäubungsbiologie, Morphologie,
Taxonomie kleinblütiger Sippen des Ophrys holosericea-Aggregates in der OstÄgäis.-
J. Eur. Orch. 56 (2-4): 219-266.
Seit vielen Jahren besuchen wir eine Reihe der ostägäischen Inseln, um die
dortigen Orchideen zu studieren. Besonderes Augenmerk legten wir für diese
Arbeit auf die verschiedenen Populationen der kleinblütigen Arten der Ophrys
holosericea-Gruppe, die sich auf jeder der besuchten Inseln andersartig
darstellten. Unter “kleinblütig“ fassen wir aus eher pragmatischen Gründen
diejenigen Formen zusammen, die eine Lippenlänge von 1 cm und weniger
haben. Um diese besser verstehen zu können und vor allem ihren taxonomischsystematischen
Status begründen und festlegen zu können, besuchten wir vor
allem die größeren Inseln Samos und Kos, daneben auch die kleinen Inseln
Phournoi, Tilos und Lipsi. Um die betreffenden Populationen gegebenenfalls
besser als Biospezies begründen zu können, studierten wir vor allem die
Bestäubungsbiologie. Folgende Ergebnisse können präsentiert werden:
1. Auf der Insel Samos beschäftigten wir uns mit Vorkommen von zwei
verschiedenen Arten der kleinblütigen Hummel-Ragwurz. Eines im Zentrum
nördlich der Stadt Mytilini mit einer stattlichen Population in einem extensiv
genutzten Weinberg. Diese Population wurde unter dem Namen Ophrys
ampelaki M.Hirth, Paulus, Kreutz bereits vorab im neuen Feldführer von
Kreutz (2024) veröffentlicht. Ihr Bestäuber ist die kleine Langhornbiene
Eucera digitata. Eine andere vor allem im Südwesten der Insel im Raum
Limnionas-Agia Kyriaki. Diese Form wird als Ophrys kerkissoula M.Hirth &
Paulus in diesem Artikel neu beschrieben. Die Art kommt auch auf der
Nachbarinsel Phournoi vor. Ihr Bestäuber ist Eucera laxiscopa. Beide Formen
unterscheiden sich in der Blütenmorphologie und haben verschiedene
Autorenexemplar - copyright by AHO Baden-Württemberg
Journal Europäischer Orchideen 56 (2-4): 2024. 221
Bestäuber.
2. Von der kleinen Insel Lipsoi wird eine kleinblütige holosericea gemeldet,
aber nicht abschließend beschrieben, da die Datenbasis unzureichend und der
Bestäuber unbekannt ist.
3. Ophrys tili von der Insel Tilos wurde erneut untersucht und mit den sehr
ähnlich aussehenden Pflanzen von Kos verglichen, die bereits früher als
O. ellinicaea beschrieben wurden. Da beide Formen kaum zu unterscheiden
sind und beide denselben Bestäuber Eucera cineraria haben, halten wir sie
für konspezifisch.
4. In einer phylogeographischen Erörterung werden die heutigen Verbreitungen
der Arten im Zusammenhang der Daten von KOUGIOUMOUTZIS et al. (2017)
der prä- und postglazialen Landveränderungen der östlichen Ägäis diskutiert.
Summary
Hirth, M. & H.F. Paulus (2024): Pollination biology, morphology, taxonomy of
the small flowered species of the Ophrys holosericea aggregate in the East
Aegean.- J. Eur. Orch. 56 (2-4): 219-266.
For many years, we have been visiting several of the East Aegean islands to
study the orchid flora there. For this work, we paid particular attention to the
different populations of the small-flowered Ophrys holosericea group, which
presented themselves differently on each of the islands visited. As smallflowered,
we group together plants of the Ophrys holosericea aggregate with lip
length less than 1 cm. To understand these better and, above all, to be able to
establish their taxonomic-systematic status, we visited the larger islands of
Samos and Kos in particular, as well as the smaller islands of Phournoi, Tilos
and Lipsi. To be able to establish the populations concerned as a biospecies, we
studied the pollination biology in particular. The following results can be
presented.
1. On the island of Samos, we discovered the presence of two different “small
flowered holosericeas”. One in the center, north of the city of Mytilene, with
a substantial population in an extensively used vineyard. The other mainly in
the southwest of the island, in the Limnionas-Agia Kyriaki area. Both forms
differ in flower morphology and have different pollinators. The vineyard
population has already been published in advance in the new field guide by
KREUTZ (2024) under the name Ophrys ampelaki M.Hirth, Paulus, Kreutz. Its
pollinator is the small long-horned bee Eucera digitata. The other form is
Autorenexemplar - copyright by AHO Baden-Württemberg
220 Journal Europäischer Orchideen 56 (2-4): 2024.
described as O. kerkissoula M.Hirth & Paulus in this article. This species also
occurs on the neighbouring island of Phournoi. Its pollinator is Eucera
laxiscopa.
2. From the small island of Lipsoi. a small flowered holosericea is reported but
not finaly discribed, as the data base is insufficient and the pollinator is
unknown.
3. Ophrys tili from the island of Tilos was re-examined and compared with the
very similar-looking plants from Kos which had already been described as
O. ellinicaea Kreutz & B. Tenschert (2011). Since both forms are hardly to
distinguish and both have the same pollinator, Eucera cineraria, we consider
them to be conspecific.
4. In a phylogeographic context, the current distributions of the species are
discussed with regard to data presented by KOUGIOUMOUTZIS et al. (2017) on
pre- and postglacial land changes in the eastern Aegean.
... maritima appear to be identical from a cytological point of view. According to Greuter (1979), HCL: Haploid total length, TF: Total form percentage (Huziwara 1962), AsI: Karyotype asymmtry index (Arano and Saito 1980), CV CL : Coefficient of variation of chromosome length, CV CI : Relative variation in centromeric index, AI: Asymmetry index (Paszko 2006), A 1 : Intrachromosomal asymmetry, A 2 : Interchromosomal asymmetry (Romero Zarco 1986), M CA : Mean centromeric asymmetry (Peruzzi and Eroğlu 2013), S Cl: Stebbins classification (Stebbins 1971). ...
The chromosome number and karyomorphology of two subspecies of the Anthemis maritima complex collected from two different coastal localities in the Skikda region (northeastern Algeria) are reported in this study: Anthemis maritima subsp. maritima, which is common throughout the Mediterranean, and A. maritima subsp. bolosii, a strict Algerian endemic recently rediscovered after 100 years of disappearance. The Feulgen staining method indicated that Anthemis maritima subsp. maritima is a tetraploid with 2n=4x=36 chromosomes (2n=11m+3sm+4st), and Anthemis maritima subsp. bolosii is a diploid with 2n=2x=18 chromosomes (2n=4m+3sm+2st). Both taxa have symmetrical karyotypes, 2A and 1A, respectively, according to Stebbins classification. These findings are novel for both the subspecies Anthemis maritima subsp. bolosii and the Algerian population of Anthemis maritima subsp. maritima.
... The geographic distribution of taxa and genetic lineages in the Balkans, Aegean Archipelago, and Anatolia document their historic range dynamics. The Aegean Sea poses an effective barrier to Balkan-Anatolian (European-Asiatic) dispersal for many plant species (e.g., Greuter 1979;Runemark 1980;Strid 1996). This is manifested as the phytogeographical 'Rechinger's line' (cf. ...
The Balkan Peninsula is widely acknowledged as one of the centers of European biodiversity and a major glacial refugium for plant species. Its extensive geographic heterogeneity and diverse mosaic of habitats, coupled with relatively high environmental stability over long periods, have enabled the diversification of lineages and simultaneously fostered the long-term survival of diverse species. The increasing number of phylogeographically studied plant species in the region has substantially contributed to a better understanding of the relationships among closely related taxa and provides significant insights into intraspecific lineage differentiation. The present paper reviews the current knowledge on plant phylogeography in the Balkan Peninsula. By providing an updated overview of the most recent studies dealing with the diversity and evolution of Balkan plants, we explore the phylogeographic patterns and roles of refugia in structuring genetic diversity and highlight the crucial evolutionary processes that shaped the diversity of plants in the region. Molecular clock-based estimations highlight the importance of Pleistocene climatic fluctuations across taxonomic groups and lineage distribution patterns corroborate the persistence of multiple glacial refugia. Spatial congruence in phylogeographic splits is determined and discussed. An examination of phylogeographic connections with adjacent regions (i.e., the Alps, Apennine Peninsula, Asia Minor, Carpathians, and central Europe) uncovers several consistent patterns. Additionally, allopatric and ecological speciation, polyploidy, and hybridization are identified as crucial evolutionary mechanisms acting in the Balkan Peninsula and shaping species diversity. Furthermore, the existing research gaps are identified and future approaches with the potential to better understand the unique Balkan flora are highlighted.
... (d'après GREUTER 1979, modifié). tains interglaciaires, les transgressions marines ont parfois haussé le niveau de la mer jusqu'à 35 m au-dessus du niveau actuel, anéantissant les végétaux des parties basses des îles (GREUTER 1970(GREUTER , 1971(GREUTER , 1979. 60 2009 encore, de nombreux complexes touristiques sont en cours de construction dans ces zones. ...
... This is in line with previous works stating that climate has also contributed to the biogeographical configuration of the Aegean archipelago (Kougioumoutzis et al. 2014a(Kougioumoutzis et al. , 2017Kougioumoutzis and Tiniakou 2014). This is also in harmony with the fact that nearly 40% of the present Aegean flora (excluding the endemics) has reached the Aegean Islands owing to human action in prehistoric or early historic times (Greuter 1979). ...
The Aegean archipelago has long been the main research area of numerous biogeographers, plant ecologists and taxonomists due to its intricate palaeogeography and high environmental and topographical heterogeneity. Nevertheless, some parts of this archipelago are essentially unexplored and the processes driving spatial variation in species composition remain unaddressed. Aiming to fill these gaps, we investigated the flora and plant diversity patterns of the Northern Sporades island group, as well as its biogeographical relationships. The study area lies in the biogeographical region of the West Aegean islands and comprises 23 islands and islets. The total flora of the study area consists of 1202 infrageneric taxa, belonging to 517 genera and 120 families, reflecting its geographical and bioclimatic characteristics. The endemic element consists of 41 taxa (3.4% of the flora), eight of which are restricted to the West Aegean islands and two are single island endemics. Area emerged as the most important variable in shaping plant species richness, while niche-based processes played a lesser role in driving these patterns. Regarding the taxonomic and phylogenetic beta-diversity patterns, environmental filtering and not dispersal limitation seems to shape the plant assemblages of the Northern Sporades islets. Biogeographically, the Northern Sporades island group seems to be closer connected to the Kiklades rather than to Evvia or the adjacent mainland, due to their longer isolation and separate palaeogeographical history during the Quaternary.
... εηώλ έλαληη πεξίπνπ 10.000-21.500 εηώλ ησλ ππνινίπσλ λεζηώλ (Meulenkamp 1971, Sondaar 1971, Greuter 1979, Υξηζηνδνπιάθεο 1986, Carlström 1987, Perissoratis & Conispoliatis 2003, (β) ηεο ρισξηδηθήο ζύλδεζεο κε ηα λεζηά ηνπ Ν. Αηγαίνπ σο κέξνο ηνπ Ν. Αηγαηαθνύ Σόμνπ (Greuter 1971) θαη (γ) ηεο κεγάιεο απόζηαζεο από ηα βνξεηόηεξα κεγάια λεζηά. ...
The aim of the thesis is the taxonomic and biogeographical approach of the Labiatae family in the East Aegean (E. Aegean) focusing on Chios Island (E. Aegean Islands, Greece) and Çeşme-Karaburun Peninsula (Anatolia, Turkey). The following were carried out: (i) taxonomy of the Labiatae plants in Chios and Çeşme-Karaburun, and mapping of their distribution in the two regions (ii) study of their distribution patterns in the two regions in relation to the habitat types and geological substrates they occupy, (iii) study of the infraspecific variation of their essential oil content in Chios in relation to the habitat types and geological substrates they occupy, and (iv) study of the biogeographical affinities of Chios and Çeşme-Karaburun based on the presence or absence of Labiatae plants.
The Labiatae family is represented in Chios Island by 51 and in Çeşme-Karaburun Peninsula by 44 taxa (species and subspecies), which belong to 23 and 21 genera, respectively. These taxa belong to four subfamilies: Nepetoideae, Lamioideae, Ajugoideae and Scutellarioideae. Eleven taxa are new records for Çeşme-Karaburun, namely Ajuga orientalis, Ballota nigra subsp. ruderalis, Melissa officinalis subsp. altissima, Mentha suaveolens, Nepeta italica, Salvia pomifera subsp. calycina, S. tomentosa, Teucrium montanum, Thymbra spicata, Thymus sipyleus and Ziziphora taurica. The distribution of forty-eight Labiatae taxa was mapped in 1 km x 1 km grid.
The taxonomic approach of the Labiatae taxa in Chios and Çeşme-Karaburun showed that: (i) In the Spicatae group of the genus Mentha, sect. Mentha the co-occurrence of the allied M. longifolia, M. suaveolens subsp. suaveolens and M. spicata results possibly to gene flow among the three taxa and thus to complex hybrid populations. (ii) In the genus Micromeria the characters that can be used as diagnostic in Chios and Çeşme-Karaburun are the calyx characters: presence or absence of a ring of hairs in the throat, teeth divergence, hair type and upper calyx teeth length. (iii) In the species Teucrium divaricatum, Stachys cretica, Thymus sipyleus and Ziziphora taurica have been recorded morphological characters of different subspecies that are reported by different authors for the E. Aegean region. The morphological overlap is possibly related to the geographical position of Chios and Çeşme-Karaburun at the phytogeographical boundary between Europe and Asia. (iv) Regarding the taxa occurring in both Chios and Çeşme-Karaburun, no morphological differences were recorded in the material examined from the two regions, which is possibly related to the recent isolation of Chios from Çeşme-Karaburun, which took place ca. 11,500-21,500 years ago.
Based on their number of records in the habitat types of Chios and Çeşme-Karaburun the species were divided into seven groups: (i) species growing mostly in habitat types of the supra- and oro-Mediterranean, (ii) species growing mostly in anthropogenic and riparian habitat types, (iii) species growing mostly in anthropogenic habitat types, (iv) species without preference to specific habitat types, (v) species growing mostly in shrub-dominated habitat types of the thermo- and meso-Mediterranean, (vi) species growing mostly in arable land, and (vii) species growing mostly in pine forests (only in Çeşme-Karaburun).
Based on their number of records in the three main rock categories of Chios and Çeşme-Karaburun, Melange (ME), Mesozoic carbonate platform (ΜΑΠ) and Caenozoic-recent rocks (ΚΠ), the species were divided into five groups: (i) species growing mostly in the ΜΑΠ, (ii) species growing mostly in the ME and in ΚΠ, (iii) species growing mostly in ΚΠ, (iv) species growing mostly in the ΜΑΠ and in ΚΠ, and (v) species without preference to specific geological substrate.
According to the number and geographical position of the grid cells they occupy, the taxa were divided into five categories according to the distribution pattern they follow: common, locally common, locally restricted, sporadic and rare. The species‟ distribution patterns seem to be related to the habitat types and geological substrates they occupy. Most interesting are the distribution pattern of the locally common species in the northern part of Chios, which seems to be related to the distribution of rocks categories on the island and the distribution pattern of the rare species which seem to follow three trends: (i) rare species growing in habitat types with limited distribution in the study area, (ii) rare species growing in habitat types with large distribution in the study area (non-anthropogenic), and (iii) rare species growing in anthropogenic habitat types, mostly settlements.
The Melange seems to be rather interesting in relation to the distribution of species, especially in Chios, but also in Çeşme-Karaburun. Some species, mainly hygrophilous, show a locally dense distribution in the Melange, while other species- even common ones- seem to “avoid” it. The preference of hygrophilous species is possibly related to the different water permeability of rocks of the Melange comparing to neighbouring rocks. The “avoidance” of the Melange is possibly related to its chemical composition (presence of heavy metals).
The Aipos plateau in the central part of Chios seems to act as a barrier, with natural (abrupt increase of elevation) and anthropogenic (degradation, grazing pressure) characteristics, in the dispersal of species on the island from south to north and vice versa, contributing possibly to the geographical differentiation of the species distribution on the island.
The essential oil content of leaves and inflorescences of 13 Nepetoideae species (Salvia fruticosa, S. sclarea, Mentha suaveolens, M. longifolia, M. spicata, M. pulegium, Thymbra capitata, Th. spicata, Origanum onites, Satureja thymbra, Clinopodium insulare, Melissa officinalis and Lavandula stoechas) varies from 0.08 mL 100 g-1 d. w. (Melissa officinalis) to 6.99 mL 100 g-1 d. w. (Origanum onites). The highest essential oil content values (> 6 mL 100 g-1 d. w.) were recorded in the shrubby species Th. capitata, O. onites and Satureja thymbra. The correlation of the infraspecific variation of essential oil content to the habitat types and geological substrates showed that: (i) the majority of the shrubby species (Thymbra capitata, Origanum onites, Satureja thymbra, Lavandula stoechas) tend to have the highest values of essential oil content in shrub-dominated habitat types, (ii) in the Mentha species, which are azonic, the infraspecific variation of essential oil content does not seem to be related to the habitat type or geological substrate, and (iii) Thymbra capitata and Satureja thymbra have statistically significant differences in the essential oil content between different habitat types and/or geological substrates.
The habitat types classification, which contrary to the traditional classifications of vegetation includes geographical and abiotic environmental factors, seems to be a useful tool in the study of species distribution and the variation of their essential oils.
Chios and Çeşme-Karaburun demonstrate close biogeographical affinities, which are supported by: (i) the similar chorological and life-form spectra, (ii) the high percentage of Anatolian/Balkan taxa that occur in both regions, and (iii) the high similarity indicated by the Sørensen and Jaccard indexes and the hierarchical cluster analysis. The E. Aegean Islands seem to be a floristically heterogenic island group which shows an unclear differentiation from north to south at the dividing level of Samos and Ikaria.
The taxonomic and biogeographical study of the Labiatae plants in Chios and Çeşme-Karaburun highlights the special geographical position of the two regions as part of a transitional zone- the E. Aegean region- at the phytogeographical boundary between Europe and Asia. Although Chios and Çeşme-Karaburun belong to distinct phytogeographical regions- the E. Aegean Islands and Anatolia, respectively- this distinction seems to lie in the political borders separating the two regions and not some natural geographical boundary. The close biogeographical affinities of Chios with Çeşme-Karaburun, closer than with any other island in the E. Aegean, reflect the recent isolation of Chios from the neighbouring peninsula and advocate that the two regions form part of one single phytogeographical entity.
... beobachteten jetzt-Zustand erklärbar machen, zumal bei der von GREUTER (1979) anhand zahlreicher Beispiele postulierten, unter den Bedingungen kleinräumiger insulärer Isolation vorherrschenden Stasis. Das Beispiel von Crepis ×cytherea Kamari, einer hybridogenen, im ehemaligen Überlappungsbereich der Eltern-Areale entstandenen und dort stabilisierten hybridogenen Art (GREUTER 1979: 100), passt genau zum Muster der hier behandelten Hypericum-Sippen. ...
A new subspecies of Hypericum perfoliatum is described and named. It is endemic to some Cycladean islands (Paros, Seriphos, and Naxos). On Andros and Naxos, it forms obviously hybridogenous populations with H. perfoliatum subsp. perfoliatum, the nominal taxon that is widespread on mainland Greece. The genesis of this distributional pattern is discussed. In agreement with earlier assumptions, it is interpreted as resulting from immigration of the mainland taxon in phases of lowered sea level during Pleistocene glaciation maxima, followed by evolutionary stasis during the subsequent period of insular isolation.
... In addition, the high phylogenetic divergence between Ph. pygmaeum and the other Mediterranean taxa could suggest that this taxon is a relict element from an older Aegean or Mediterranean flora that went extinct elsewhere. In fact, some authors reached the same conclusion for the majority of the endemics of Crete, which was isolated from the mainland for 5 million years during the Pliocene (Greuter, 1979). Although within the Irano-Turanian clade the resolution is poor, the morphological and molecular evidence indicates that Ph. acuminatum and Ph. ...
The precise generic delimitation of Aliella and Phagnalon and their tribal affinities are at present unresolved. The main goals of our study were to verify the monophyly of these two genera and to determine their closest affinity group within Gnaphalieae. We analysed sequences of the trnL intron and trnL‐trnF spacer of Gnaphalieae and other Compositae tribes, in order to elucidate the tribal position of Aliella, Macowania, Phagnalon and Philyrophyllum. In addition, we analysed ribosomal nrDNA together with the ycf3‐trnS and trnT‐trnL spacers of cpDNA to elucidate the relationships within Aliella and Phagnalon. The genera Anisothrix, Athrixia and Pentatrichia are closely related to Aliella and Phagnalon. Aliella, Macowania and Phagnalon are nested within the "Relhania clade", and the subtribal affinities of Philyrophyllum lie within the "crown radiation clade". The monophyly of Aliella and Phagnalon is not supported statistically and Aliella is paraphylethic in most of the analyses. The resulting phylogeny suggests an African origin for Aliella and Phagnalon and identifies three main clades in Phagnalon, the Irano‐Turanian clade, the Mediterranean‐Macaronesian clade and the Yemenite‐Ethiopian clade. Some endemics to Yemen and Ethiopia are resolved in the Mediterranean‐Macaronesian clade, providing new evidence of phytogeographical links between Macaronesia, Eastern Africa and Southern Arabia. Incongruence between the chloroplast and nuclear molecular data and the lack of resolution in some clades may indicate that hybridization could have played an important role in the evolution and diversification of Phagnalon and Aliella.
... Since the Kiklades islands (home of H. graecum) and the Peloponnisos (H. laconicum) were connected through mainland Greece until the end of the Pliocene (c. 2 Ma ;Fattorini 2002;Chatzimanolis et al. 2003), but subsequent eustatic sea-level changes during the Pleistocene were never severe enough to reconnect the Kiklades to mainland Greece (Greuter 1979;Comes et al. 2008;Poulakakis et al. 2015 (Liveri et al. 2018) may constitute retained plesiomorphies from the joint ancestor and shows that allopatric differentiation processes through genetic drift and/or selection in the island populations of the former species may still be in full swing. ...
Hymenonema (Compositae, tribe Cichorieae) together with the genera Catananche, Gundelia, and Scolymus forms the subtribe Scolyminae. It is endemic to Greece and consists of two species, Hymenonema laconicum and Hymenonema graecum, which occur in the south Peloponnisos and central Aegean area, respectively. The present contribution aims at a phylogenetic reconstruction of evolutionary relationships among the 12 species of the subtribe, focusing on the temporal and spatial framework for its evolution. The phylogenetic relationships among the members of Scolyminae were inferred from molecular data based on the multi-copy region of the nrDNA internal transcribed spacers ITS1 and ITS2, two intergenic spacers of the cpDNA (trnL-trnF, rpl32-trnL), and one single-copy nuclear region (D10). The gene trees were reconstructed using Bayesian phylogenetic methods. All gene trees support the monophyly of Hymenonema and the sister-group relationship with the genus Scolymus. The further sister-group relationship of this group (Hymenonema–Scolymus) with Catananche is also supported by nrDNA and cpDNA analyses. Finally, a species tree (inferred in a Bayesian coalescent framework) was reconstructed and dates the divergence time between the two Hymenonema species to the Pleistocene (around 1.3 Ma ago). Maximum likelihood-based biogeographical reconstructions suggest a Miocene (pre-Messinian) differentiation of the subtribe on the northern Tethyan platform, followed by Miocene/Pliocene dispersal events to the western Mediterranean and North-African platforms and final, small-scale vicariance events within the genera in the Pleistocene.
... Leaves sheathing more than ¾ of the stem and a life cycle in which dormancy is almost absent are characters that coexist in a few other Greek Allium species belonging to three different sections, viz. A. archeotrichon and A. makrianum C. . Characters like these have been considered as biological traits and adaptations in the arid climatic conditions prevailing in the Mediterranean area during the late Tertiary, i.e. more than 5 million years ago, and the species concerned have been considered as relict floristic elements (Greuter 1979;Kollmann & al. 1990;Tzanoudakis & Kypriotakis 1993, 2008Brullo & al. 1997;Tzanoudakis 2000). ...
Allium symiacum Galanos & Tzanoud., from the island of Symi (SE Aegean, Greece), is described as a species new to science. It is an autumn-flowering, single-island endemic species of A. sect. Codonoprasum (Amaryllidaceae) and is classified as Critically Endangered according to IUCN Red List categories and criteria. Considering the morphological and karyological characters of the new species, its possible relationships to other autumnal species of A. sect. Codonoprasum distributed in the E Mediterranean area are discussed.
... This reflects insularity, the role of islands as refuges and the adaptation of species to island environments, especially aridity/moisture gradients. The flora of the Mediterranean islands can be classified into three broad categories according to their origin (Greuter 1979): i) a relict element: ancestors of pre-isolation phase, ...
... agropyroides, growing also in the Aegean "cliff flora" (cf. RUNEMARK 1971), indicates that this species might have been present in the region already before the Aegean Continent had started in the early Pliocene to become definitely an archipelago (MALDONADO 1985), a fundamental geological event which explains the otherwise enigmatic Aegean phytogeography in many other groups (EH~ENDOPd~ER 1958, GREUTER 1979. The absence of H. convolutum in the Aegean Islands is more obscure. ...
The geographical distribution of the grass genus Helictotrichon in the Mediterranean Region is analysed and mapped for 33 taxa. Based on only a single life form (perennial herb), a variety of edaphically, climatically and altitudinally differently adapted species complexes has evolved in the area. Most of these complexes show west-east disjunctions and contain geographically, sometimes even edaphically vicarious taxa with complementary distribution. A transition from mesomorphic to xeromorphic habit occurred independently in different species groups and led to the establishment of the 'modern' Mediterranean taxa which are in part highly polyploid derivatives of more mesophilic diploids. The significance of polyploidy, patterns of parapatric and sympatric distribution, biogeographical borders, and centres of species diversity are discussed in context with the history of the Mediterranean vegetation. New combinations are: Helictotrichon setaceum subsp. petzense, H. pratense subsp. lusitanicum. H. praetutianum.
... The catastrophic effects of the Messinian salinity crisis (Krijgsman et al., 1999) and the Pliocene land-plate movements that formed the Peloponnese and Aegean Islands (Popov et al., 2006;Ivanov et al., 2011) also had a great influence *Corresponding author. E-mail: borata@man.poznan.pl on contemporary geographical ranges of species (Greuter, 1979;Thompson, 2005). ...
Juniperus drupacea is an eastern Mediterranean mountain tree with a disjunct geographical range. We hypothesized that this disjunct occurrence (the Peloponnese in Europe and the Taurus and Lebanon Mountains in Asia) should be reflected in the patterns of genetic and morphological diversity and differentiation. Nuclear microsatellite markers (nSSR) and biometric variables of the cones and seeds were examined on material sampled from four populations in Europe and eight in Asia. The Asian populations were characterized by a higher level of genetic diversity than the European populations. The genetic differentiation among populations was moderate but significant (FST = 0.101, P < 0.001). According to the clustering performed with BAPS, six genetically and geographically groups of populations were found: I and II from the Peloponnese; III from the Taurus Mountains; IV and V from the Anti-Taurus Mountains; and VI from the Lebanon Mountains. The level of genetic differentiation among these six groups (4.30%, P = 0.012) probably reflects long-lasting genetic isolation during the Pleistocene, as limited genetic admixture was found. In accordance with genetic analysis, the biometric investigations indicated a high level of morphological divergence between the European and Asian populations of the species, with further differentiation between the populations from the Taurus and Lebanon Mountains.
... In fact, the most suggestive evidence for the just mentioned so-called nonadaptive radiation comes from some plant groups or some beetle families (mainly Tenebrionidae) inhabiting the ecologically fairly homogenous continental island system of the Aegean Archipelago (e.g. GREUTER 1969, COMES et al. 2008FATTORINI 2002 andCHATZIMANOLIS et al. 2003 for Tenebrionidae). SCHLÜTER et al (2007SCHLÜTER et al ( , 2009SCHLÜTER et al ( , 2011 ...
... Rare species in anthropogenic habitats. The human factor has been determinant in shaping the Aegean flora (Cellinese et al., 2009;Greuter, 1979;Triantis & Mylonas, 2009) with proportions of the anthropogenic element reaching up to one third or even almost half of some ...
The East Aegean (East Aegean Islands, Greece and West Anatolia, Turkey) is a biogeographically transitional region, where biodiversity elements from Europe and Asia join. However, affinities in the region were until recently scarcely explored. We assess biogeographical affinities in the East Aegean focusing on distribution patterns of Lamiaceae plants in Chios Island and its adjacent Çeşme–Karaburun Peninsula. Detailed in-situ record was acquired for 48 native species. These were grouped based on their habitat and geological substrate preference, their distribution was mapped in grid cells and distribution patterns were analysed in relation to species groupings. In both Chios and Çeşme–Karaburun, species follow five distribution patterns: widespread, locally widespread, locally restricted, sporadic and rare. Fifty to 62% of the species exhibit similar distribution patterns, trends in habitat and geological substrate preference in Chios and Çeşme–Karaburun, results complying with previous evidence of close biogeographical affinities of the East Aegean Islands and neighbouring Anatolia. Differences observed between the two regions may be attributed to insularity effects, human impact and the melange, an old rock matrix known for its key role in elucidating regional geodynamic evolution. Distributions of widespread and locally widespread species in Chios give evidence of density compensation and niche shifts responses, however, the actual occurrence of these phenomena in island plant populations is still to be elucidated. Overall, the species distribution patterns, particularly those of rare ones, reflect the complex geological history, palaeogeography and human influence in the Aegean region.
... Although a long-standing tradition exists in the study of Mediterranean plant biodiversity and endemism (e.g. Contandriopoulos & Favarger, 1962, 1974Pignatti, 1978;Greuter, 1979Greuter, , 1991Dahlgren, 1991;Petit & Thompson, 1999, and references therein) only in the past few decades has increased attention been paid to the tempo and mode of the present-day distribution of Mediterranean plants, with special attention to taxa with narrow geographical ranges and often small populations. These studies have greatly benefited from the advent of simple methods for retrieving DNA-based information, and, in the recent past, numerous well-founded studies investigating the phylogenetic relationships in connection to the biogeography of a range of plant taxa, or even entire clades, have been published (e.g. ...
The present investigation investigated the genetic structure of a monophyletic group of endemic species belonging to the Genista ephedroides species group: G. bocchierii, G. cilentina, G. demarcoi, G. dorycnifolia, G. ephedroides, G. gasparrinii, G. insularis, G. numidica, G. tyrrhena subsp. tyrrhena, G. tyrrhena subsp. pontiana and G. valsecchiae, all distributed in the western Mediterranean. Using seven plastid microsatellites, 16 populations (288 individuals) were screened. Haplotype fixation was observed in particular for most of the Tyrrhenian taxa (i.e. G. bocchierii, G. cilentina, G. demarcoi, G. ephedroides, G. gasparrinii, G. insularis, G. tyrrhena subsp. tyrrhena and G. valsecchiae). However, although genetic diversity within populations was low [(hS = 0.132 (± 0.056)], a high level of total plastid DNA diversity [hT = 0.866 (± 0.042)] was detected, and analysis of molecular variance indicated that variation is almost exclusively expressed among populations (95.25%). The plastid microsatellites identify two groups of taxa, one including Sardinian taxa and populations of G. tyrrhena subsp. pontiana and the other including two subgroups, one of which includes Sicilian/Aeolian elements and the other with G. numidica/G. cilentina and G. dorycnifolia. Results allow us to consider G. cilentina as originating by recent anthropogenic dispersal and G. tyrrhena subsp. pontiana as a possible stabilized hybrid between local plants and members of the Sardinian group contributing the maternal lineage. The evolutionary history of the group possibly results from the effects of ancient events fostering geodispersal and range contraction, followed by more recent long-range dispersal or geodispersion over Pleistocenic land bridges.
... Although a long-standing tradition exists in the study of Mediterranean plant biodiversity and endemism (e.g. Contandriopoulos & Favarger, 1962, 1974Pignatti, 1978;Greuter, 1979Greuter, , 1991Dahlgren, 1991;Petit & Thompson, 1999, and references therein) only in the past few decades has increased attention been paid to the tempo and mode of the present-day distribution of Mediterranean plants, with special attention to taxa with narrow geographical ranges and often small populations. These studies have greatly benefited from the advent of simple methods for retrieving DNA-based information, and, in the recent past, numerous well-founded studies investigating the phylogenetic relationships in connection to the biogeography of a range of plant taxa, or even entire clades, have been published (e.g. ...
The present investigation investigated the genetic structure of a monophyletic group of endemic species belonging to the Genista ephedroides species group: G. bocchierii, G. cilentina, G. demarcoi, G. dorycnifolia, G. ephedroides, G. gasparrinii, G. insularis, G. numidica, G. tyrrhena subsp. tyrrhena, G. tyrrhena subsp. pontiana and G. valsecchiae, all distributed in the western Mediterranean. Using seven plastid microsatellites, 16 populations (288 individuals) were screened. Haplotype fixation was observed in particular for most of the Tyrrhenian taxa (i.e. G. bocchierii, G. cilentina, G. demarcoi, G. ephedroides, G. gasparrinii, G. insularis, G. tyrrhena subsp. tyrrhena and G. valsecchiae). However, although genetic diversity within populations was low [(hS = 0.132 (± 0.056)], a high level of total plastid DNA diversity [hT = 0.866 (± 0.042)] was detected, and analysis of molecular variance indicated that variation is almost exclusively expressed among populations (95.25%). The plastid microsatellites identify two groups of taxa, one including Sardinian taxa and populations of G. tyrrhena subsp. pontiana and the other including two subgroups, one of which includes Sicilian/Aeolian elements and the other with G. numidica/G. cilentina and G. dorycnifolia. Results allow us to consider G. cilentina as originating by recent anthropogenic dispersal and G. tyrrhena subsp. pontiana as a possible stabilized hybrid between local plants and members of the Sardinian group contributing the maternal lineage. The evolutionary history of the group possibly results from the effects of ancient events fostering geodispersal and range contraction, followed by more recent long-range dispersal or geodispersion over Pleistocenic land bridges.
... Permanence in situ and prolonged evolutionary standstill are salient characteristics of Mediterranean island floras (Greuter, 1979). These have a relictual nature and are at least ancient as the islands themselves; for Cyprus, Crete and the Baleares about 5-6 milion years, dating back to the post-Messinian transgression (Greuter, 1995). ...
Characteristics of Mediterranean island floras are analyzed with stress on endemic units. On these bases the main relationships between the major Mediterranean areas and the inland ter-ritories with the strongest floristic affinities are analyzed. Finally the role of aliens in Mediter-ranean island floras and threats are discussed. st International Congress "Insularity and Biodiversity", May 11 th -13 th , 2012 -Palermo (Italy) INTRODUCTION The Mediterranean is among the richest regions in the world for wild and cultivated species. Cir-cum-Mediterranean countries house about 25,000 species, almost one tenth of the world's vascular flora, the 63% of which are endemic (Greuter, 1991; Medail & Quezel, 1999). A peculiarity of this area is the high amount of species with a narrow range, many of which are local endemics. In some sectors endemics are more than 20%. Among these there is Sicily and the other major Mediterranean islands and island groups: Baleares, Corse, Sardinia, Kriti, Cyprus with their mountain areas. The main reason for this high endemism can be searched in the pronounced habitat fragmentation that characterises the Mediterranean area as a whole. CHARACTERISTICS OF MEDITERRA-NEAN ISLAND FLORAS Mediterranean island flora is relatively well known although each year new species, sometimes completely unknown, are described (e.g. Ptiloste-mon greuteri Raimondo et Domina. Fig.1). Tradi-tionally the floras of large Mediterranean islands are considered ancient, relatively poor in species, rich in endemic taxa and particularly vulnerable (Greuter, 1995). Permanence in situ and prolonged evolutionary standstill are salient characteristics of Mediterra-nean island floras (Greuter, 1979). These have a re-lictual nature and are at least ancient as the islands themselves; for Cyprus, Crete and the Baleares about 5-6 milion years, dating back to the post-Mes-sinian transgression (Greuter, 1995). According to the equilibrium theory of island biogeography (MacArthur & Wilson, 1967) the number of species on an island is in a state of dynamic equilibrium; diversity eventually stabilizes but turnover remains high as species continuously colonize and go ex-tinct. The flora of the Mediterranean was subject to an impoverishment due to the climatic fluctuation during Pliocene and a subsequent enrichment due to immigration by long range dispersal (Quézel & Médail, 2003). This is more manifest in the islands where original species pool seems to become
... Between 350 and 250 ka, the subaerial land was extended, the sea was restricted and almost 50-60% of the present Aegean Sea was land with extensive drainage systems, delta plains and large lakes (Lykousis, 2009). Owing to late Pleistocene sealevel regressions, the gaps between mainland Greece and the Cyclades were reduced in width (Greuter, 1979). During the Last Glacial Maximum (LGM; c. 20 ka), the area between Evvia, Attiki and Kea was partly exposed to subaerial conditions, while several palaeolakes existed within its central part (Lykousis, 2009). ...
AimAlthough the factors shaping plant species richness patterns across the islands of the central Aegean are well known, the processes driving the assembly of these island communities remain unclear. To shed light on these processes, we identified biogeographical modules within the phytogeographical area of the Cyclades and tested for nestedness across the islands.LocationThe Cyclades, Greece.Methods
We used a network approach to detect island biogeographical roles and modules, based on a large and detailed database of the Greek endemic plant taxa of the Cyclades, and we tested for nestedness in the island–species matrices.ResultsThe Cyclades were significantly modular and divided into five biogeographical modules. Three of the modules were significantly nested and two displayed all four possible biogeographical roles (connectors, module hubs, network hubs, peripherals). Most of the network's taxa are classified as peripherals and widespread endemics.Main conclusionsThe borders of the five modules correspond remarkably well to the palaeogeographical and climatic compartmentalization of the Cyclades. The flora of the Cyclades has not yet reached the relaxation phase and the region may act as an ecogeographical filter for the distribution of several plant lineages. Naxos, Milos and Anafi play an important role for the network's connectivity, while at least five adjacent phytogeographical regions affect the distribution patterns of the endemic taxa present in the Cyclades.
... In Gestalt eines Nachschlagewerkes entsteht eine "Ökotafel" ("Logarithmentafel für Pflanzenökologen". Ellenberg 1974, 1979. Aber nicht nur Einzelarten an sich werden so standörtlich klassifiziert, sondern es besteht zudem die Möglichkeit, Pflanzengruppen, komplette Pflanzengemeinschaften oder Pflanzengesellschaften zu bewerten. ...
... Collins et al. (2012) have also attributed the changes in the composition and structure of landscapes in the Mediterranean islands to anthropogenic activity, as well as to the climate changes that occurred during the late Holocene. According to Greuter (1979Greuter ( , 1995Greuter ( , 2001, nearly 45% of the present Aegean flora reached the Aegean islands as the result of human action in prehistoric or early historic times; a large part of the Aegean flora is synanthropic (occurring in man-made habitats or in habitats affected by livestock) and the proportion of such species in the Aegean increases with grazing (Bergmeier and Dimopoulos 2003). Greuter (1995) also noted that these synanthropic species may be perfectly integrated into native plant communities. ...
Background: The South Aegean Volcanic Arc (SAVA), one of the most notable geological structures of the Mediterranean Sea, is floristically well-known. Nevertheless, the factors that contribute to shaping the plant species richness of the SAVA remain unclear.
Aims: Investigate factors that affect plant species richness and identify plant diversity hotspots in the SAVA and other central Aegean Islands.
Methods: We used stepwise multiple regressions to test the relationship between several environmental factors and plant species richness in the SAVA, as well as the residuals from the species-area linear regressions of native, Greek and Cycladian endemic taxa as indicators of relative species richness.
Results: Area was confirmed as the most powerful single explanatory variable of island species richness, while geodiversity, maximum elevation and mean annual precipitation explained a large proportion of variance for almost all the species richness metrics. Anafi, Amorgos and Folegandros were found to be endemic plant diversity hotspots.
Conclusions: We demonstrated that geodiversity was an important factor in shaping plant species diversity in the Cyclades, while mean annual precipitation, human population density and maximum elevation were significant predictors of the Greek endemics present in the Cyclades. Finally, Anafi was found to be a plant diversity hotspot in the South Aegean Sea.
... East Anatolia and the Transcaucasus are the primary centres of origin of the genus Centaurea while the Mediterranean area and the Balcan Peninsula are secondary centres (Routsi and Georgiadis, 1999;Wagenitz, 1975;Wagenitz and Hellwig, 1996). The Mediterranean Basin is considered a refugium for many of these species (Greuter, 1979) and several, endemic taxa of Centaurea are very narrowly distributed in the region (Pisanu et al., 2011). Within the genus Centaurea, the group of C. cineraria (Cela-Renzoni and Viegi, 1982;Pignatti, 1982;) comprises ten species (Hilpold et al., 2011). ...
A case study on Centaurea gymnocarpa Moris & De Not., a narrow endemic species, was carried out by analyzing its morphological, anatomical, and physiological traits in response to natural habitat stress factors under Mediterranean climate conditions. The results underline that the species is particularly adapted to the environment where it naturally grows. At the plant level, the above-ground/below-ground dry mass (1.73 ± 0.60) shows its investment predominately in the above-ground structure with a resulting total leaf area per plant of 1399 ± 94 cm2. The senescent attached leaves at the base of the plant contribute to limit leaf transpiration by shading soil around the plant. Moreover, the dense C. gymnocarpa leaf pubescence, leaf rolling, the relatively high leaf mass area (LMA = 12.3 ± 1.3 mg cm−2) and leaf tissue density (LTD = 427 ± 44 mg cm−3) contribute to limit leaf transpiration, also postponing leaf death under dry conditions. At the physiological level, a relatively low respiration/photosynthesis ratio (R/PN) in spring results from high R [2.26 ± 0.59 μmol (CO2) m−2 s−1] and PN [12.3 ± 1.5 μmol (CO2) m−2 s−1]. The high photosynthetic nitrogen use efficiency [PNUE = 15.5 ± 0.4 μmol (CO2) g−1 (N) s−1] shows the large amount of nitrogen (N) invested in the photosynthetic machinery of new leaves, associated to a high chlorophyll content (Chl = 35 ± 5 SPAD units). On the contrary, the highest R/PN ratio (1.75 ± 0.19) in summer is due to a significant PN decrease and increase of R in response to drought. The low PNUE [1.5 ± 0.2 μmol (CO2) g−1 (N) s−1] in this season is indicative of a greater N investment in leaf cell walls which may contribute to limit transpiration. On the contrary, the low R/PN ratio (0.05 ± 0.02) in winter is resulting from the limited enzyme activity of the respiratory apparatus [R = 0.23 ± 0.08 μmol (CO2) m−2 s−1] while the low PNUE [3.5 ± 0.2 μmol (CO2) g−1 (N) s−1] suggests that low temperatures additionally limit plant production. The experiment of the imposed water stress confirms that the C. gymnocarpa growth capability is in conformity with the severe conditions of its natural habitat, likewise as it may be the case with others narrow endemic species that have occupied niches with similar extreme conditions.
... agropyroides, growing also in the Aegean "cliff flora" (cf. RUNEMARK 1971), indicates that this species might have been present in the region already before the Aegean Continent had started in the early Pliocene to become definitely an archipelago (MALDONADO 1985), a fundamental geological event which explains the otherwise enigmatic Aegean phytogeography in many other groups (EH~ENDOPd~ER 1958, GREUTER 1979. The absence of H. convolutum in the Aegean Islands is more obscure. ...
The geographical distribution of the grass genusHelictotrichon in the Mediterranean Region is analysed and mapped for 33 taxa. Based on only a single life form (perennial herb), a variety of edaphically, climatically and altitudinally differently adapted species complexes has evolved in the area. Most of these complexes show west-east disjunctions and contain geographically, sometimes even edaphically vicarious taxa with complementary distribution. A transition from mesomorphic to xeromorphic habit occurred independently in different species groups and led to the establishment of the “modern” Mediterranean taxa which are in part highly polyploid derivatives of more mesophilic diploids. The significance of polyploidy, patterns of parapatric and sympatric distribution, biogeographical borders, and centres of species diversity are discussed in context with the history of the Mediterranean vegetation. New combinations are:Helictotrichon setaceum subsp.petzense, H. pratense subsp.lusitanicum, H. praetutianum.
... Nigella; Bittkau & Comes, 2005) and roughly coincides with the well established floristic Rechinger's line (Strid, 1996). The existence of this barrier to plant migration and gene flow appears to stem from the palaeogeographical evolution of the region through the Miocene and early Pliocene (Greuter, 1979; see also Bittkau & Comes, 2005, and references therein). Indeed, our phylogeographical reconstruction (Fig. 3a) agrees with an ancient split and posterior differentiation of the H1 and H2 lineages in the Balkan-Aegean and easternmost Mediterranean domains, respectively. ...
Aim The post-glacial range dynamics of many European plant species have been widely investigated, but information rapidly diminishes as one moves further back in time. Here we infer the historical range shifts of Laurus, a paradigmatic tree of the Tethyan flora that has covered southern Eurasia since the Oligo-Miocene, by means of phylogenetic and phylogeographical analyses.
... The close affinities of the Labiatae in the two regions reflect the recent isolation of Chios from the neighbouring peninsula, which took place during the Pleistocene, ca. 20.000 years ago [45]. Further analysis taking into account the distribution patterns of the Labiatae plants in the E. Aegean, in relation to the geological evolution of the Aegean region, might reveal new data on the phytogeographic status of the family in the E. Aegean region. ...
This article presents a bilateral cooperation between School of Biology, Aristotle University of Thessaloniki (Greece) and Department of Biology, Çanakkale Onsekiz Mart University (Turkey), under the auspices of the European Lifelong Learning Programme-Erasmus. An Erasmus placement grant was provided to a doctoral student of Aristotle University, for the study of the affinities of Labiatae plants between Chios Island [East Aegean Islands (EAI), Greece] and the adjacent Çeşme-Karaburun Peninsula (Anatolia, Turkey). The study resulted in close affinities of the Labiatae in Chios and Çeşme-Karaburun [high similarity indices, similar chorological spectra, and occurrence of narrowly distributed species (Anatolia-EAI-Balkan endemics) in both regions]. Additionally, extensive field work diroughout Çeşme-Karaburun Peninsula gave 11 new records for this region, meeting the first objective of the European Plant Conservation Strategy regarding documentation of plant diversity, in Anatolia, one of the biodiversity hot-spots of the Mediterranean basin. The article concludes with the benefits for a doctoral student through such crossborder mobility cooperations and further attempts to think towards a common concept in the East Aegean, where bom the East Aegean Islands and the adjacent Anatolian mainland will be studied as one phytogeographic entity. This cooperation is the first mutual step of Greece and Turkey for a joint study on the phytogeography of the East Aegean, a region, where political borders are of no phytogeographic sense.
... In the Mediterranean region, islands are for the most part fragments of land that have become isolated due to their separation from continental areas; the few exceptions include the islands originated from submarine volcanic activity, such as the Aeolian Islands. The former are usually known as "continental islands", the latter "oceanic islands", even if a different terminology was proposed by Greuter (1979): namely, "chersogenous" for the former, "thalassogenous" for the latter ones. ...
The present paper briefly provides the state of the art of the knowledge on vascular plant endemism in the oceanic (“thalassogenous”) Aeolian Archipelago (Sicily). Preliminary analysis of distribution areas and review of recent literature on biosystematics of endemic species revealed that: (a) Aeolian strictly endemic taxa are just 6, i.e. about the 0.7 % of the local vascular flora; among them, just 4 can be considered (with doubt) derived from in situ evolution. (b) The other 18 endemics are taxa occurring in wider areas, so they cannot be generally considered “locally evolved” but relicts. This preliminary analysis confirms that not only continental (“chersogenous”) but all Mediterranean islands are primarily conservative rather than evolutionary active systems.
Anthropogenic climate and land use change pose major threats to island floras worldwide, yet few studies have integrated these drivers in a single vulnerability assessment. Here, we examine the endemic flora of Evvia, the second-largest Aegean island in Greece and an important biodiversity hotspot, as a model system to address how these disturbances may reshape species distributions, community composition, and phylogenetic diversity patterns. We used species distribution models under the Ensemble of Small Models and the ENphylo framework, specifically designed to overcome parameter uncertainty in rare species with inherently limited occurrence records. By integrating climate projections and dynamic land use data, we forecasted potential range shifts, habitat fragmentation, and biodiversity patterns for 114 endemic taxa through the year 2100. We addressed transferability uncertainty, a key challenge in projecting distributions under novel conditions, using the Shape framework extrapolation analysis, thus ensuring robust model projections. Our findings reveal pronounced projected range contractions and increased habitat fragmentation for all studied taxa, with more severe impacts on single-island endemics. Our models demonstrated high concordance with established IUCN Red List assessments, validating their ecological relevance despite the sample size limitations of single-island endemics. Current biodiversity hotspots, primarily located in mountainous regions, are expected to shift towards lowland areas, probably becoming extinction hotspots due to projected species losses, especially for Evvia’s single-island endemics. Emerging hotspot analysis identified new biodiversity centres in lowland zones, while high-altitude areas showed sporadic hotspot patterns. Temporal beta diversity analysis indicated higher species turnover of distantly related taxa at higher elevations, with closely related species clustering at lower altitudes. This pattern suggests a homogenisation of plant communities in lowland areas. The assessment of protected area effectiveness revealed that while 94.6% of current biodiversity hotspots are within protected zones, this coverage is projected to decline by 2100. Our analysis identified conservation gaps, highlighting areas requiring urgent protection to preserve future biodiversity. Our study reveals valuable information regarding the vulnerability of island endemic floras to global change, offering a framework applicable to other insular systems. Our findings demonstrate that adaptive conservation strategies should account for projected biodiversity shifts and serve as a warning for other insular biodiversity hotspots, urging immediate actions to maintain the unique evolutionary heritage of islands.
Simple Summary
Aiming to cope with the provisions of Aichi Biodiversity Targets, EU Biodiversity Strategy and EU Green Deal, we conducted the first nationwide, phylogenetically informed identification of vascular plant diversity hotspots and endemism centres in Greece. By this, we identified the most important factors that shaped them, and assessed the effectiveness of the Special Areas of Conservation of the Natura 2000 network in safeguarding them. Qualitative and quantitative results are provided and presented in thematic maps and relevant diagrams, highlighting areas of conservation importance, and identifying current protection scheme gaps. Simultaneously, our work contributes to national efforts for drafting Natura 2000 sites Management Plans, as well as to the MAES implementation in Greece.
Abstract
Biodiversity hotspots (BH) cover a small fraction of the Earth’s surface, yet host numerous endemics. Human-induced biodiversity loss has been increasing worldwide, despite attempts to halt the extinction crisis. There is thus an urgent need to efficiently allocate the available conservation funds in an optimised conservation prioritization scheme. Identifying BH and endemism centres (EC) is therefore a valuable tool in conservation prioritization and planning. Even though Greece is one of the most plant species-rich European countries, few studies have dealt with the identification of BH or EC and none has ever incorporated phylogenetic information or extended to the national scale. Consequently, we are unaware of the extent that Special Areas of Conservation (SAC) of the Natura 2000 network efficiently protect Greek plant diversity. Here, we located for the first time at a national scale and in a phylogenetic framework, the areas serving as BH and EC, and assessed the effectiveness of the Greek SAC in safeguarding them. BH and EC are mainly located near mountainous areas, and in areas supposedly floristically impoverished, such as the central Aegean islands. A critical re-assessment of the Greek SAC might be needed to minimize the extinction risk of the Greek endemics, by focusing the conservation efforts also on the BH and EC that fall outside the established Greek SAC.
Cliffs are among the environments with the most adverse conditions for living organisms because of the limited availability of soil, moisture, and nutrients, and owing to the harsh conditions of exposure. They constitute shelters for rare, endemic, and range-restricted plant taxa. A main database has been prepared which includes vascular plant taxa that are obligate or facultative chasmophytes and also contains information about their life form, chorology, protection status, occurrence in more than 135 places such as cliffs, gorges, or open rocky habitats, and their geographical distribution in the 13 phytogeographical regions of Greece based on available floristic, vegetation, and phytosociological literature and on the authors' own collections and observations. The paper presents an analysis of the total diversity of cliff plant species, as well as the diversity of obligate chasmophytic plant species, endemics, and range-restricted taxa, in addition to the results of studying the distribution patterns of different subsets of plant taxa in the different phytogeographical regions of Greece. Hemicryptophytes and chamaephytes are the dominant life forms of the chasmophytic taxa. Among 935 species and subspecies registered, 476 are obligate chasmophytes, of which the majority are Greek endemics. Hierarchical cluster analysis of different subsets of plant taxa revealed affinities of the cliff flora of different phytogeographical regions. Additionally, 15 chasmophytic taxa mentioned in Annexes II, IV, and V of EEC Directive 92/43 belong to the cliff flora, of which 10 are obligate chasmophytes and nine have a priority for protection. Eighteen taxa are included in the IUCN Red List of Threatened Species, of which four are critically endangered (CR), seven are vulnerable (VU), and three are endangered (EN). The distinct correlation between endemism and chasmophytic ecology is emphasized, since a detailed understanding of the local distribution and specific habitats of rare plants provides an opportunity for local conservation efforts that can influence biodiversity conservation on a larger scale. © 2017 Institute of Botany and Botanical Garden Jevremovac, Belgrade.
The Achillea ageratifolia and A. clavennae groups (Asteraceae, tribe Anthemideae) were investigated for intra- and interspecific leaf flavonoid variation. Numerous flavonoid glycosides, mostly unidentified, and variously methylated free aglycones based on scutellarein, 6-hydroxyluteolin, quercetagetin, and 6-hydroxy-kaempferol were detected. The distribution of methylated aglycones among individuals of A. ageratifolia was used to model the putative biosynthetic pathways. Several steps in the biosynthesis probably involve enzymes (methyltransferases) with strict positional specificities and broader substrate specificities. Geographical patterns in the flavonoid content of A. ageratifolia were only partly consistent with the current concept of A. ageratifolia. Different flavonoid profiles in A. ageratifolia do not appear to be of crucial importance in its adaptation to different habitats. The species of the A. clavennae group were clearly separable on flavonoid profiles and the presence of a hybrid population was supported by the flavonoid data. Interspecific differences in number of glycosides in relation to the number of free aglycones are discussed.
Islands frequently have distinctive and often unique assemblages of species. In general they have lower species diversity than equivalent continental areas, but tend to have elevated numbers of endemic species. The number of species in a particular taxonomic group on a given island and the proportion of these which are endemic appears to depend on a wide variety of factors, both historical and ecological. Among these are the degree of isolation, age, size, topography and climate of the island and the biological characteristics of the taxonomic groups concerned, in particular their vagility (the ease with which they disperse). Historical accident also appears to play a large part in patterns of species occurrence on islands.
The development of plant population biology (SOLBRIG et al., 1979) as a discipline which connects plant systematics with ecology has been one of the most important trends within recent years, though at present more questions have been asked than answered. Likewise, the study of reproductive biology, notably of tropical plants, has turned into a field of tremendous activity; most of it will be reviewed in this volume under pollination ecology, but the results are of course of great interest for the evolution and systematics of seed plants. Methodological progress has been made in the interpretation of phytochemical data. Apart from that, the trends mentioned in the last review (Progr. Bot. 41, 239) are still of interest. As to the problems of classification in general, the number of cladistic studies has increased, but it is still doubtful if they will have a major impact. Comprehensive, integrated approaches to the systematics and biology of major groups include the Solanaceae symposium (HAWKES et al., 1979), and the resumption of the Natürliche Pflanzenfamilien series with a multi-authored treatment of the Loganiaceae (LEEUWENBERG, 1980).
Chromosome counts and the results of anatomical investigation are reported for 9 species of the genus Helictotrichon in the western and southern Mediterranean. Chromosome numbers of 3 rare species are reported for the first time: 2n = 126 in H. lusitanicum (N Portugal, endemic), 2n = 28 in H. jahandiezii (Moyen Atlas, endemic), and 2n = 105, 122-124 in H. pruinosum (NW Africa). Ploidy levels of 6x (2n = 42) in the W Mediterranean species H. marginatum and H. albinerve are new. Morphological and anatomical characters of H. lusitanicum and a peculiar Moroccan Atlasic variant of H. pruinosum ('subvar. dolosa') are reported in detail and discussed with chromosomal, biogeographical and taxonomic aspects concerning the evolution of Helictotrichon subg. Pratavenastrum in the Holarctic flora. General phylogenetic problems referring to 'reticulate evolution' and 'retention' of primitive character states in the high polyploids of Helictotrichon are exemplified. Aims of future research in Helictotrichon and further Aveneae are outlined. The name H. lusitanicum is a new combination, denoting a new endemic species of the Portuguese flora.
In classic studies of island biota three main approaches may be identified: description of strange plants and animals as phenomena at the species level; examination of the mechanisms behind evolutionary and distributional patterns recognized at the population and community level; and quantitative and comparative evaluation of the diversity of entire island biota.
The variation and evolution of reproductive traits in island plants have received much attention from conservation and evolutionary biologists. However, plants on islands in the Mediterranean region have received very little attention. In the present study, we examine the floral biology and mating system ofCyclamen creticum, a diploid perennial herb endemic to Crete and Karpathos. Our purpose is to quantify (1) variation and covariation of floral traits related to the mating system, (2) the ability of the species to self in the absence of pollinators and its relative performance on selfing and outcrossing and (3) genetic diversity within and among populations. Pollen/ovule ratios were indicative of a xenogamous species. A controlled pollination experiment showed that the species is self-compatible but is unable to set seed in the absence of pollinators, probably due to stigma-anther separation. A multiplicative estimate of inbreeding depression based on fruit maturation, seed number and percentage seed germination gave δ=0.38. Population genetic diversity was high, 54.76% polymorphic loci, a mean of 1.78 alleles per locus and a mean observed heterozygosity of 0.053.F-statistics nevertheless indicated high inbreeding rates (meanFis=0.748) in natural populations, and low levels of population differentiation (meanFst=0.168).C. creticumthus appears to have a mixed-mating system with high levels of (pollinator) mediated inbreeding (either by facilitated selfing, geitonogamy or biparental inbreeding) in natural populations.
An analysis is made of the floristic richness of the Sierra Nevada (southern Spain), where more than 2100 vascular plants were catalogued, thus making it the area with the richest and most varied flora in the western Mediterranean region, including a considerable number of endemic species. In a study of the endangered flora and its protection, 116 taxa are listed as having the most restricted distribution and as being under threat of extinction. Field work for these taxa was undertaken in order to improve knowledge of their habitats, populations, chorology and main threats. Using the new IUCN categories of threat, an analysis was made for each of the taxa, especially those that are critically endangered with a very low number of individuals or very narrow areas of occupancy. Mention is made of the problems posed by the practical application of the new IUCN categories. The main types of environmental impact affecting the flora are also described, including grazing, natural causes, wetland desiccation and water pollution. The highest-lying areas (oro- and cryoro-Mediterranean belts) are where most of the threatened plants are to be found. Since these same plants are mostly endemic to the Sierra Nevada, the need to protect the high peaks of the area is paramount. Reference is made to current legislation protecting threatened plants and the gaps in the European Union Habitats Directive with relevance to the protection of the Sierra Nevada flora. A number of key strategies for protection are proposed.
The variation and evolution of reproductive traits in island plants have received much attention from conservation and evolutionary biologists. However, plants on islands in the Mediterranean region have received very little attention. In the present study, we examine the floral biology and mating system ofCyclamen creticum, a diploid perennial herb endemic to Crete and Karpathos. Our purpose is to quantify (1) variation and covariation of floral traits related to the mating system, (2) the ability of the species to self in the absence of pollinators and its relative performance on selfing and outcrossing and (3) genetic diversity within and among populations. Pollen/ovule ratios were indicative of a xenogamous species. A controlled pollination experiment showed that the species is self-compatible but is unable to set seed in the absence of pollinators, probably due to stigma-anther separation. A multiplicative estimate of inbreeding depression based on fruit maturation, seed number and percentage seed germination gave δ=0.38. Population genetic diversity was high, 54.76% polymorphic loci, a mean of 1.78 alleles per locus and a mean observed heterozygosity of 0.053.F-statistics nevertheless indicated high inbreeding rates (meanFis=0.748) in natural populations, and low levels of population differentiation (meanFst=0.168).C. creticumthus appears to have a mixed-mating system with high levels of (pollinator) mediated inbreeding (either by facilitated selfing, geitonogamy or biparental inbreeding) in natural populations.
Zusammenfassung Funktionelle Vielfalt ist Vielfalt, die darauf ausgerichtet ist, zum Pflanzenschutz ge- gen Krankheiten, Schadinsekten und Unkräuter und abiotische Stressfaktoren beizu- tragen. Sowohl mechanistische Interaktionen, wie Verdünnungs- und Barrierenef- fekte als auch komplexe ökologische und pflanzenphysiologische Prozesse, wie Al- lelopathie und induzierte Resistenz können in diversifizierten Beständen wirken. Am Beispiel einer Untersuchung des traditionellen Reisanbausystems in Bhutan wird aufgezeigt, dass ein wichtiger zusätzlicher Prozess, der in vielfältigen Systemen stattfindet, die gegenseitige Anpassung der Kulturpflanzen und der Pathogene (d.h. Ko-Evolution) ist. Ko-Evolution hat in Bhutan dazu geführt, dass in Regionen, wo der Krankheitsdruck durch Pyricularia grisea hoch ist, wirksamere und vielfältigere Resi- stenzen vielfältigeren Virulenzen gegenüber stehen als in Zonen, wo der Krankheits- druck geringer ist. In herkömmlichen Systemen wird Vielfalt durch Sorten- und Ar- tenmischungen, verkleinerte Schlaggrößen und Fruchtfolgen erzielt. Es kommen ge- netisch einheitliche Sorten zum Einsatz. Ko-Evolution ist nur möglich, wenn Variation in den Kulturpflanzen für die zu selektierende Eigenschaft (z.B. Resistenz) vorhan- den ist. Um in der Landwirtschaft Ko-Evolution zu fördern, sind Züchtungsansätze gefordert, die Diversität für Resistenz produzieren. Der ökologische Landbau wird sich in Zukunft noch verstärkt die wichtigen Funktionen der Biodiversität im Pflanzen- schutz zu eigen machen müssen, um Erfolg zu haben und stellt so einen wichtigen Partner für innovative Ansätze in der Pflanzenzucht dar. Summary Functional diversity is diversity that is designed to contribute to plant protection against diseases, insect pests, weeds and abiotic stress. Mechanistic interactions such as dilution and barrier effects as well as complex ecological and plant physio- logical processes, such as allelopathy and induced resistance contribute to plant
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