PhD-excursion to Crete 3-10 October 2009

Transcription

1 PhD-ecursion to Crete 3-10 October 2009 Editor: Daniel Udd 1 Authors: Gustaf Granath 1, Camille Madec 1, Adriana Puentes 1, Kate St.Onge 2, Daniel Udd 1 and Maria Uscka-Perzanowska 1 Supervisor: Francesco Spada 1 1 Plant Ecology, Department of Ecology and Evolution, EBC, Uppsala University, Norbyvägen 18D, Uppsala, Sweden 2 Evolutionary Functional Genomics, Department of Evolution, Genomics and Systematics, Uppsala University, Norbyvägen 18D, Uppsala, Sweden

2 Front picture: Spiny cusion of Sarcopoterium spinosum in the phrygana. Photo:DU Photographers in the report: Daniel Udd (DU) and Camille Madec (CM)

3 Summary Crete is an ecellent place to study plant geography. We had the chance to see remnants from the last glaciation period, but also plants introduced by humans. The spectacular landscape also gave us the possibility to study different environments and how they shape plant communities. We could also see an overall zonation from sea level up to the highest peaks, but the flora was surprisingly similar along this climatic gradient. The Cretan landscape is not, in our opinion, severely degraded or over-used as the Mediterranean landscape sometimes is described as. Undoubtedly, grazing and human eploration have had an impact, but there still remain many forested areas where the different abiotic conditions are less stressful. Hence, abiotic factors should be considered in relation to anthropogenic factors when the impact of these is being assessed. The Cretan flora also contains many fascinating adaptations to dry conditions and grazing. We were able to see how plants changed morphologically due to grazing or humidity; at times, it was hard to understand that two completely different individuals were actually members of the same species. Furthermore, to have the opportunity to see wild growing pistachio, for eample, and other eotic and culturally important plants was one of the many highlights of our trip. 3

4 Table of Contents Introduction...6 People...6 Landscape...6 Climate...7 Wildlife...7 Vegetation types...8 Woodlands...9 Maquis...9 Garigue-Phrygana...9 Steppe...9 Above the Timberline Cliffs Seashores Wetlands Cultivated areas Diary Day 0: Arrival Day 1: Elafonisi Day 2: Up the mountains

5 Day 3: Heading for the summit, Lefka Ori Day 4: Imbros Gorge Day 5: Date Palm Beach Day 6: Knossos Day 7: Back to Chania Acknowledgements Appendi A: Species list 5

6 Introduction The Mediterranean Island of Crete was the subject of study for the SVS (Svenska Vätgeografiska Sällskapet) ecursion 3-10 October This Greek island hosts many spiny cushions, pygmy trees, gorges and goats, which shape the unique Cretan landscape. Because of the islands proimity to several major landmasses and the history of the rising and falling of the Mediterranean Sea, the islands flora is of particular phytogeographical interest. People Humans first settled the island in Neolithic times, bringing a variety of animals and plants. Several sites on the island are of great historical significance, for eample Knossos, which is a site of the Minoans, Europe s oldest civilization. Crete was the center of this ancient civilization, which established itself on several Mediterranean islands. Iraklion, today s capital, is the centre of modern Crete with a population of over 300,000. Chania is the second largest city with approimately 70,000 inhabitants. Fishing and farming have traditionally been the main occupation of the people of Crete. However, a recent shift towards the tourist industry has made a major change in Crete s economy. Today Crete is one of the most popular sunny destinations in Europe with over 2 million tourists reaching the island every year. Landscape Crete is the largest Greek island and the second largest in the eastern Mediterranean. The islands most striking geographical feature is the numerous gorges, which cut through the three mountain ranges: the White Mountains or Lefka Ori (2,452 m), the Idi Range (2,456 m) and the Dikti Mountains (2,148 m). These gorges where formed by uneven up heaving of the islands rock, creating jagged cracks, which are slowly worn smooth by erosion. The rough walls and crumbled rocky floor of these gorges are home to many endemic plants. We visited the Imbros gorge and hiked western slopes of the Samaria gorge to one of the summits of Lefka Ori (1,960 m). There are many fertile mountain plains among these mountains. These plains collect the water runoff from the mountains and with it silt and debris. The typical soil is deep brown-yellow overlying or mied with gravel, ideal for cultivation. We visited the mountain plain of Omalos. 6

7 The 1046 kilometers of beautiful coastline which bring tourist to Crete also creates a special niche for many plants. Crete is surrounded by four seas: to the north is the Sea of Crete, in the east the Karpathion Sea, to south is the Libyan Sea and the Myrtoan Sea is to the west. We have visited several beaches, most notable Elafonisi. Climate Winter is the wet season on Crete, with a flourishing plant life. Plants have to withstand severe drought during the hot, dry summer. Some of them respond to this drought with growth reduction or growth cessation, some of them (annuals) die back. July is the hottest month with an average high of 28 C, Jan-Feb is the coldest with an average low of 8 C. The summer months are truly hot and dry with an average precipitation of 0 mm. During the wet season precipitation varies widely across the island, the largest difference is seen between high and low altitudes. It has been estimated that precipitation on Lefka Ori could reach 2000 mm of rain and snowfall in a year. At the other end of the spectrum are the low coastal regions, the south-east coast is epected to be the driest with an estimated 240 mm of rain per year. Wind is also a formidable force on Crete. Wind-storms are known to whip through the mountains affecting the morphology of trees, for eample creating flag shaped cypress. Wildlife Crete, similar to other large Mediterranean islands, was home to a variety of interesting mammals, which are now etinct: pygmy elephants and hippopotami, giant rodents and insectivores, and several species of deer roamed the island in the late Pleistocene or early Holocene. Most fossils of these animals were collected from caves, which are difficult to date. Today the only large animals roaming the island are the feral and domesticated goats. The goats were most likely first brought to the island with the Minoans. These goats play a large part in shaping the flora of Crete. Their grazing induces interesting morphologies in several tree species. Although Crete lacks diverse mammals, many interesting bird species are found there. Of particular interest are the large vultures and griffons, including the bearded vulture Gypaetus barbatus. 7

8 Vegetation types The particular history of Crete makes its flora quite unique! The island is roughly equidistant from Europe, Asia and Africa but was connected to them 23 million years ago, when the Sea dried up, allowing flora and fauna to migrate. The latest connection with mainland dates back to late Miocene (the Messinian period, 6 to 5 Million years ago). Thus, nowadays, floristic elements from the three continents co-eist in Crete. Moreover, the Cretan flora contains a high degree of endemic species (10%), which can be eplained by the eistence of high mountains. These functioned as isolated botanical islands together with the fact that Crete has been isolated from the mainland for about 5 million years. A profound change in the Cretan vegetation (variation in plant community structure and composition) since the Neolithic has been partly attributed to the climate having become more arid, but also by the activities of man and his grazing animals. During the first settlements about 8000 years ago, the climate was more humid but not wet enough for continuous forests. The Mediterranean climate that Crete eperiences today was fully developed years ago. Much of the vegetation today shows obvious adaptation to the Mediterranean climate, such as small leaves with thick surface and/or densely covered with hairs (features that reduce transpiration during the dry season), as well as adaptation to grazing, such as presence of spines and/or a pungent taste. A common misconception about Crete, and the Mediterranean landscape in general, is the so called massive environmental degradation caused by earlier human societies. It is said that magnificent forests used to cover these areas but men cut down the forests and let the goats graze so no regeneration was possible and the ground became bare. However, vegetation can be affected by several other factors and climate is probably the most important one. There is little doubt that human activities have played a big part in shaping the landscape, but other factors such as climate, are usually ecluded from the story on how the Mediterranean landscape was formed. The fact that vegetation has re-grown in abandoned areas (ceased cultivations, reduced grazing) during periods with favourable climatic conditions (wetter), illustrates that the landscape is not ruined and is affected by many factors. The devastating effects of goat grazing should also be put into perspective. Crete was once populated by several grazing mammals and intense browsing should be viewed as a natural state to which flora plant communities adapted during a long time span. This means that at some etent an open landscape is probably the aboriginal structure of the 8

9 local vegetation. Here we describe the different vegetation zones today present in Crete. Woodlands The aboriginal woodland of Crete, which covered much of the area, was degraded and reduced due to woodcutting, browsing and burning. Nowadays, Cretan forest covers a very small part (<5 %) of the island. It consists mostly of Mediterranean sclerophylls, such as Acer sempervirens, Cupressus sempervirens, Phillyrea latifolia, Pistacia terebinthus, Quercus coccifera and Q. ile. On the richer soils, some nonsclerophyllous species, like Quercus pubescens, can be found. African accent of Cretan forests gives coastal populations of an endemic palm Phoeni theophrasti. From the other hand, forests of the endemic tree Zelkova abelicea are an Asiatic element of Cretan mountain massifs. Maquis In general, the term maquis represent Mediterranean shrub communities. Most Shrubs in Crete are trees reduced to a shrubby form, whereas undershrubs are woody plants which can never grow into trees. Maquis is often nearly continuous in west Crete, with patches of phrygana or steppe. Garigue-Phrygana Phrygana is a Greek term, first used by Theophrastus, describing small, tough-leaved thorny shrubs and dwarf shrubs. This vegetation type is typical for Mediterranean ecosystems with severe browsing and wildfires. The characteristic grey-green patches of phrygana can be found from sea-level up to the mountain zone. Phrygana consist mostly of spiny, cushion-shaped, low shrubs, resistant to drought and grazing. Under the protection of thorny shrubs there may be patches of multitude geophytes, including several orchids. Nowadays communities composed of spiny undershrubs also are called garigue, and many authors use this term and its Greek equivalent interchangeably. Steppe Steppe is the name for communities of herbaceous plants; it occurs where the land is too dry and/or where the ground is too compact for the development of woodland and other communities. The vegetation is similar to North Africa and South-western Asia. 9

10 Steppe vegetation is more predominant in east Crete, with varying amounts of phrygana, where the climate is hotter and drier. Above the Timberline The natural limit of woodland in Crete etends from about 1450m on the northern mountain slopes to m in sheltered southern slopes. Northerly winds during the growing season probably prevent trees from reaching higher altitudes on the northern side. The alpine and arctic-alpine elements of the central and northern Spiny cusion-vegetation in a hillside phrygana. Photo:DU European mountains (such as Alpine grassland) are absent, replaced by a miture of Balkan and Asiatic orophytic elements. Above the timberline, the landscape is mainly composed of rocks (limestone or dolomite) and often appears devoid of vegetation. Cliffs Chasmophytes are plants typical for cliffs because of their ability to tolerate the harsh conditions of vertical surfaces, such as etreme temperatures, drought effect and weak 10

11 soil layer. These highly specialized plants are usually effectively restricted to cliff habitat due to the lack of competitiveness. As a consequence several endemic species of chasmophytes can be found on Cretan cliffs: Petromarula pinnata, Hypericum jovis (central Crete), Hypericum amblycaly (eastern Crete), Hypericum aciferum (southern coast of western Crete). Seashores Some maritime areas support only shrub communities because storms and salt spray have prevented the formation of woodland. These could be the origin of many of the phrygana plants. The marine communities of sandy beaches of Crete share many species in common with the coasts of the rest of Europe. There are also some islets surrounding the Cretan coasts, which have a distinct flora due to dry rocky conditions and low grazing intensity. As cliff communities (described earlier), few chasmophytes occur on these islets because of tectonic movements and fluctuations in sea level. Wetlands Wetland habitats in Crete regroup few natural or artificial lakes, rivers, marshes and streams, where widespread European species as Iris pseudacorus, Phragmites australis and Vite agnus-castus occur. Some endemic species are also present, such as Lathyrus neurolobus net to rivers and streams, as well as Care idaea and Cirsium creticum in marshes. Cultivated areas The most fundamental and characteristic aspect of Cretan agriculture is the cultivation of olive tree with its basic products: olive oil and olive fruits (main ingredients of the Cretan gastronomy). The olive tree, Olea europea, has been grown in Crete since Minoan times and nowadays cultivation constitutes a half of the total cultivated area of the island. Thus, the grey-green stretches of olive groves are the typical components of the Cretan landscape. 11

12 Diary Day 0: Arrival We arrived late in the evening at the airport outside Chania, got the cars and headed for the place where we were going to stay for the two first nights. The hotel was in the touristic old harbor. Day 1: Elafonisi After the first night in Chania we headed out west along the coast. At our first stop, which was just outside Chania, we could see scattered tamarisk trees Tamari cretica on the well-grazed beach pastures, like a savannah. We continued uphill where the roads were sided by olive trees. In the valleys the vegetation was richer with several species of oak (e.g. Quercus coccifera) and the big broad leafed Platanus orientalis, a tree that we would see lots of during our trip. We continued going west and stopped for lunch in a small village. Cretan salad was everyone s favourite here and the fresh vegetables were delicious. Our net goal was Elafonisi. We had to cross the mountain range to get there. On our way there we saw our first gorge, with goats climbing up and down the steep sides. It was obvious that not even such inaccessible areas were spared from grazing. In the village of Elos and its surroundings we saw sweet chestnut trees Castanea sativa along the road. However, it seems to be debated if the tree is indigenous or was brought to the island by humans. Eventually we reached our main attraction for this day, Elafonisi, with its lagoon bounded by islets and sandbars that you can wade out to. Although the area has been eploited for tourism, most of the coast line is still rather intact. This is a Natura2000 area and there were signs apologising that some parts had been fenced off to keep the tourists out. Interestingly, the fenced areas were irrigated. We wondered if the species they are trying to save (a drought, salt and grazing tolerant species) will be outcompeted due to this treatment. Along the coast there is a pygmy forest, mainly constituted by maritime juniper Juniperus oycedrus ssp. macrocarpa, but also some Tamari cretica and Pinus brutia, some of them impressively big. After the narrow strip of pygmy forest the spiny shrub vegetation, phrygana, took over. This was more or less the same phrygana that we would see and investigate more the net day, but then high up in the mountains. Before we left, a few of us took a swim in the rather warm water (~23 C). 12

13 Natura2000 protection in Elafonisi, Crete. The vegetation is enclosed and the endemic plants are irrigated. Photo:DU. On our way back we stopped just outside Elafonisi and looked at the Pinus brutia forest. The area was covered, but with occuring gaps here and there, by pines creeping above the ground (ma 3-4 m high). Francesco tried to make the parallel to a savannah landscape if there had been sufficient grazing. After this we went back to Chania rapidly and had dinner at tourist restaurant. Unfortunately, the food was not particularly good here and we tried to avoid these kinds of restaurants during the rest of our trip. Day 2: Up the mountains We departed early from Chania to find our way south towards the famous White Mountains (Lefka Ori) of Crete. On our way there we had the chance to appreciate clearly the different vegetation belts/types that characterize the island. Once we had left the city behind us, the unmistakable olive trees Olea europaea began to appear, a widespread species in the island s landscape. As we went through the different villages we were fascinated by the many lemon Citrus lemon, orange Citrus sinensis, fig Ficus carica, and pomegranate Punica granatum trees that we saw growing in both 13

14 private gardens and cultivars. We had not yet climbed too far up the mountain road that leads to the Omalos valley, when we took our first long stop. Here we got the chance to observe important features of the human-disturbed landscape of Crete. Remnant and modern man-made terraces, for growing different cereal crops and trees (olive trees in most cases), were abundant along the slopes of the mountains. A peculiarly abundant shrubby thicket Erica manipuliflora and a spiny shrub-like tree Quercus coccifera were also quite evident along the slopes. We learned that these species could easily dominate the landscape if they were not regularly burned in programmed forest fires. We also observed artificially planted pines Pinus spp. along the cliffs. As we continued our ascent up the mountain we also observed wild-growing pear Pyrus amygdaliformis and almond trees Prunus amygdaliformis, but we stopped to observe in detail the wonderful Quercus pubescences. It s a semi-deciduous oak, with its leaves remaining green up to 1.5 years. We eamined its fruits and learned that they contain lower amounts of tannin than acorns of other oaks, yet this tree is usually preferred by grazing animals over Quercus ile. Cultivated hillsides along the road up the Lefka Ori. Photo:DU. 14

15 A browsed Quercus ile with really spiny leaves in the phrygana vegetation. Photo:DU On the following stop, we really took the time to get to know the two predominant vegetation types that occur in the island. We had the chance to observe, feel and eamine the typical species that characterize the phrygana and the maquis. Some of the most common phryganic species we observed (and felt very painfully!) were Sarcopoterium spinosum and Thymus capitatus. These species dominate most of the landscape now due to persistent human disturbance; we learned that long ago these were, most likely, broad-leafed tree dominated forests. One of the most striking Maquis species we observed was Q. coccifera. This species presents a dimorphism of the leaves due to intense grazing by feral goats and sheep, which are widespread all over Crete. When grazed, the leaves of Q. coccifera become very small, etremely spiny and grow very dense. If the plant is able to escape herbivory, the tree is able to grow tall and strong, with normal smoother leaves. The difference between the two leaf morphs is so striking, that many a times we thought of them as two different species. Both Phrygana and Maquis species occur together in Crete, with some species being classified as part of the both groups; at this site we also saw Euphorbia characias and Verbascum macurum. Moreover, we learned that if disturbance/grazing were to diminish or disappear so would many of the Phryganic species, since the 15

16 shrub-like trees of the Maquis would have the right conditions to grow to their normal size. After a few hours back on the road, we finally reached the alpine belt. Here we observed and felt the very spiny Verbascum spinosum and Berberis cretica, among many other tragacanthic (low-growing, thorny shrubs native to mountainous regions) species. Due to their spiny nature, these shrubs are able to thrive in these dry environments. In the alpine forest belt we were able to admire the magnificent cypress Cupressus sempervirens forests. The understory was rather empty, yet we managed to spot some T. capitatus, Trifolium cretica and some Daphne sericea and some Prunus prostata trees among the cypresses. Of course, after reaching the top, we began our lovely descent into the magnificent Omalos valley. There we found the small and charming village of Neos Omalos and we settled into our hotel after a long and eciting day. Day 3: Heading for the summit, Lefka Ori After a good night s sleep, we were ready to climb towards the summit of Lefka Ori Mountain and observe the changes in vegetation with the altitude. At the beginning of our ecursion (at the site of Xyloskalo) we have seen a tree of Zelkova abelicea. Later on, half way to the summit, we saw, on distant slopes, a little forest of Zelkova abelicea, apparently above the Cupressus tree-line or among it. It is probably the remnant of a deciduous forest belt of Mediterranean Mountains. Today it is a real outsider in the present climate, it is outcompeted by the more erophytic Cupressus. Zelkova abelicea is an endemic tree species of Crete, occurring only on calcareous soil. It was also present in central Europe during the quaternary and disappeared years ago. Today we can find it only in Crete. Along with a small population of a distinct species, Zelkova sicula in Sicily, it is the only representative of this Asiatic genus in Europe. Above the tree line, a rocky landscape is predominant, some places devoid of vegetation and some others with very few herbs, shrubs and small trees. During our ecursion, some of the species we observed are restricted to high mountains habitat (Acantholimon androsaceum and Colchicum cretense - were growing where the snow was standing during the winter). Others have a bipolar distribution, e.g. Sarcopoterium spinosum and Verbascum spinosum, and can be found in similar phrygana 16

17 communities at the highest altitudes as well as at sea level. We also saw two species of spiny shrubs, Astragalus creticus and A. dolinicolus, representative of the mountain phrygana. An important thing we could observe was that the altitudinal sequence of different vegetation types which in Crete is different from what we commonly see in the Mediterranean countries (Southern Europe, Northern Africa and the Middle East (table 1)). In Crete, we do not observe Cedar forests in the altitudinal stratification. The reason for this remains unclear. It could have disappeared or never have been there. We could also observe that in areas where spiny oaks and Acer co-eist, goats prefer to feed on oaks. However, up slope, where only Acer individuals are present, goats are forced to feed on them. Table 1.Altidudal gradients of the mountain vegetation from high to low altitude. The Cretan Mountains The Mediterranean, Lebanon and Atlas Mountains Spiny (tragacanthic)dwarf-shrubs Juniperus oycedrus woodland (with Berberis cretica) Cupressus sempervirens forests (with Acer sempervirens) Pinus brutia forests Quercus pubescens (semideciduous) Quercus ile Spiny (tragacanthic)dwarf-shrubs Juniperus oycedrus woodland Cedrus forests (with Abies) Quercus sp.pl. (semideciduous) with relic of Cupressus sempervirens in the upper part. Quercus ile Quercus coccifera Quercus coccifera 17

18 One part of the epedition at a summit (1,960 m) in the Lefka Ori after a several hours hike up the mountain. Photo: DU. Day 4: Imbros Gorge After an early but great breakfast we started in Omalos, crossing the plain created by collapsed caves formed by karstification of the limestone bedrock. The Omalos plain is surrounded by a lot of caves, crevasses and holes formed by dissolving of the underlying limestone bedrock by rainwater. Our first stop for the day was a natural pear-tree with cultivated forms grafted on it. The scar from the grafting was clearly visible. It is common to graft twigs from domesticated trees to old natural stems of fruit trees, sometimes even across species and families. At the mountain slopes the cypress-woods where dominating. We went further downhill through landscapes of phrygana and maquis heading for Chania. From 18

19 A grafted pear-tree in the Omalos Plain. Probably a cultivated form on a wild one. The scar is clearly visible. Photo: DU. Chania we went on to the Imbros Gorge and started our four hour walk down the canyon to the shore of the Mediterranean Sea. The gorge was formed in the Pleistocene by erosion of cracks in the bedrock. The erosion is mostly made up of streaming water and changes in temperature. Today the gorge is flooded during periods of heavy rainfall and at the time of our visit there was no water visible. The cliffs work like refuges for endemic species. All niches in the gorge are occupied, so it is very hard for alien species to invade. Along the hillsides, where the degree of disturbance is highest, Phlomis fructicosa is dominating. In the bottom of the gorge there is a transition from Acer, via Quercus, to Cupressus as the dominating species. At the cliffs, Quercus ile, Hedera heli and Ruscus aculeatus are forming a typical plant community for gorges in the Mediterranean. Some other species are also growing at the cliffs, e.g. Centaurea idaea, Arisarum vulgare, and Petromarula pinnata The common ivy Hedera heli is a shrub-like climbing plant that grows on the cliffs. There is a morphologic difference between the leaves on the juvenile sterile and adult fertile parts of the plant. The juvenile leafs are five lobed and the adult leaves are rhombic in shape. An interesting species is the maple Acer sempervirens, which can be seen in the gorge in various forms from small shrubs to full grown trees. The leaves of Acer sempervirens are usually lobed but when it is heavily browsed it gets round and small leafs at the surface of the dense cushion-like shrub that is formed. Inside the shrub the 19

20 leaves are still lobed but smaller than leaves on unbrowsed individuals. Browsed individuals also form thorns in the branch ends, as a defense against browsing. At lower altitude, Cupressus sempervirens was more dominating in the gorge. Closer to the coast the bush Vite agnus-castus became more common. This species together with Platanus orientalis are good indicators that we are close to the coast. Towards the end of the gorge, probably uncultivated wild forms of olive Olea europaea were growing scattered in the cypress woods at the slopes. After the walk we headed for the small south-coast town Chora Sfakion for some good food and relaing in the relatively warm night under the star clear sky. Imbros Gorge with low prygana vegetation together with a large hollow oak in the bottom and trees and shrubs on the cliffs. Photo: DU. 20

21 Day 5: Date Palm Beach During this ecursion we travelled along the south seashore of Crete. The main aim was to see one of the only European sites with wild growing date palms. But first on the agenda was to visit the remains of a medieval garrison. On our way along the coast we could observe very typical vegetation for the Mediterranean shore, shrubs of Vite agnus-castus frequently accompanied by Nerium oleander. We also observed the typical characteristics of the Cretan landscape in the form of maquis and phrygana vegetation. We visited the remains of the garrison called Frangokastello, i.e. the Castle of the Franks. The fort was built in the year 1371 on the seaside to protect the area from pirate incursions. The endemic Phoeni theophrasti growing by an outlet of a river at the south coast. Photo: DU. Behind the fort we saw the Ceratonia siliqua tree. Formerly this species formed the first belt in the olive forests, but nowadays these forests are mostly destroyed or domesticated and transformed into olive groves. On the beach, close to the garrison remains, we found mostly plants adapted to desert conditions like: Anagyris foetida, which loosed its leaves in the summer; Tribulus terrestris; Crithumum maritimum; Salsola kali; Euphorbia paralias; or Pancratium maritimum, flowering in the summer. There were also widely distributed perennial grasses: Phragmites australis, Hyparrhenia hirta, and perennial cane Arundo dona. Finally we reached a costal population of the endemic palm Phoeni theophrasti, which reminds of an African landscape. The forest was dense and maze-like, formed by the high, multi-branched palms and Tamari africana trees. Phoeni theophrasti is a beautiful tree, endemic to Crete, now classified as threatened. It resembles Phoeni 21

22 Restored Minoan painting in Knossos. Are the flowers some kind of Minoan kurbits? Photo: DU. dactylifera, but its fruits are inedible. The undergrowth of the palm forest consisted mainly of Ceratonia siliqua shrubs. Closer to the river there were Sali fragilis, Phragmites australis and Pistacia lentiscus growing. The presence of Platanus orientalis and spiny Rubuscus aculeatus could be interpreted as a sign of the net step in the succession of this unique habitat. Day 6: Knossos In the morning we went to see the ruins of Knossos. By the ruins we could see two species of Pinus and we tried to eamine them, which was quite hard from a distance. Elsewhere was the restoration of the buildings really impressive. Francesco guided us trough the Minoan culture, which was not appreciated by the local guides. After the visit at Knossos the group split into two groups, some of us going for a swim in the sea and the others going for a coffee in a genuine bar up in the mountains. We found what we were looking for: a bar in a small village with locals chatting in the shade served by an old woman. This was a very relaed place. The way back to Iraklion resulted in an adventure on dirt roads in the Cretan countryside, before we had convinced ourselves to go back the easier regular road. Day 7: Back to Chania On the last day we went from Iraklion back to Chania and the airport. Along the way we had a lunch stop in a small village by the sea, Georgioupoli. We relaed on the beach for a while and some of us made a small ecursion to see the beach vegetation. A very interesting discovery was the spiny and creeping form of Phragmites australis with short stiff leaves. Walking bare-foot was painful, since a lot of spiny seeds were hiding in the sand. One of the bar owners told us that she was going 22

23 A spiny form of Phragmites australis on the beach in Georgioupoli. The sand is hiding spiny fruits. Photo: DU. to put up here own small nature reserve by her bar to protect the interesting vegetation. Back in Chania we went shopping, or at least tried to. We visited the Old Harbour for the last time before we jumped into the cars for a last ride on the Cretan roads to the airport. We are really happy and glad that we have had the opportunity to do this ecursion. We have learnt a lot about the Mediterranean landscape, vegetation, history and culture, especially about Crete. This ecursion will last in our minds for a long time and hopefully we will have use of all new knowledge in our future research. 23

24 Acknowledgements We would like to thank Svenska Vätgeografiska Sällskapet for financial aid in this ecursion. We would also thank Francesco Spada for, with great knowledge, guiding us through the landscape and history of Crete. Professor Francesco Spada is teaching us about the pygmy forests on the shores of Elafonisi. Photo: CM. 24

25 Appendi A: Species list Appendi A: Species list List of species noted during trip to Crete 3-10 October Day 1 refers to 4 October. Species Day 1 Day 2 Day 3 Day 4 Day 5 Day 6 Day 7 Acantholimon androsaceum Acer sempervirens Adianthum capillus-veneris Aisnam vulgare Anagyris foetida Anchusa cespitosa Arbutus unedo Arundo dona Arusaum vulgare Asparagus aphyllus Asperula rigida Asphodelus ramosus Asplenium adianthum-negrum Astragalus angustifolius Astragalus dolinicolus Berberis cretica Bromus tomentellus Carlina sp. Castanea sativa Catapodium rigidium Centaurea idaea Ceratonia siliqua Cinocurus echinatus Cistus creticus Citrus lemon Citrus sinensis Clematis cirrhosa Colchicum cretense Crataegus sp. Crithumum maritimum Cupressus jetisimus Cupressus pyramidalis Cyclamen graecum/hederifolium 25

26 Appendi A: Species list Species Day 1 Day 2 Day 3 Day 4 Day 5 Day 6 Day 7 Cynodon dactylon Daphne oleoides Daphne sericea Daucus carota Ebenus cretica Erica arborea Erica manipuliflora Euphorbia characias Euphorbia paralias Euphorbia sp. Ficus carica Genista acanthoclada Hedera heli Hyparrhenia hirta Juniperus oycedrus Juniperus oycedrus ssp. macrocarpa Marrubium vulgare Melica rectiflora Microwedi graecica Morus alba Nerium oleander Olea europaea Origanum dictamnus Pancratium maritimum Petrohagia dianthoides Petromarula pinnata Petronella sp. Phagnalon rupestre Phleum sp. Phlomis fructicosa Phoeni dactylifera Phoeni theophrasti Phragmites australis Picnomon acarna Pinus brutia Pinus halepensis Pinus pinea Pistacia lentiscus Pistacia terebinthus 26

27 Appendi A: Species list Species Day 1 Day 2 Day 3 Day 4 Day 5 Day 6 Day 7 Platanus orientalis Prunus amygdaliformis Prunus prostrata Pteridium aquilinum Ptilostemon chamaepeuce Punica granatum Pyrus amygdaliformis Quercus coccifera Quercus ile Quercus ithaburensis Quercus macrolepis Quercus pubescens Ruscus aculeatus Saicicornia perennis Sali fragilis Salsola kali Sarcopoterium spinosum Scilla autumnalis Sclerophylla sp. Smila aspera Stachys spinosa Staehelina arborea Stipa bromoides Styra officinalis Tamari africana Tamari cretica Tamari sp. Thymelaea hirsuta Thymelaea tartonraira Thymus capitatus Tribulus terestris Trifolium cretica Triticum vilosum Urgine maritima Verbascum arcturus Verbascum macurum Verbascum sp. Verbascum spinosum Vite agnus-castus Zelkova abelicea 27

28 Vätekologi Avd. f. Ekologi och Evolution Evolutionsbiologiskt centrum Uppsala universitet Norbyvägen 18 D Uppsala