Mediterranean ecosystems Distribution Climate Soils Vegetation types Ecophysiology

Mediterranean ecosystems ¶Distribution ¶Climate ¶Soils ¶Vegetation types ¶Ecophysiology ¶Fire ecology ¶The “Ruined Landscape” hypothesis

Mediterranean landscape I

Mediterranean landscape II

Mediterranean and mediterraneoid ecosystems Only 2% of land area but ~20% of plant species; 30° - 35° latitude; west side of continents

Plant communities, El Torcal, Andalucia, Spain (photos: Ian Hutchinson, June, 2007)

Mediterranean climates Mean ann. range (°C) 10° 16° 19° July H H H

Mediterranean climates January H H H

Mediterranean Basin precipitation Feb. July mm/day Source: CLIVAR Africa

Mediterranean soils (terra rossa) A Thin mull humus forming in eroded remains of Bt horizon Btf Illuvial horizon enriched with clay and iron, has become red-coloured. High clay content renders them relatively impermeable and prone to erosion C Parent material (commonly calcareous) but in this case composed of Palaeozoic shales

Mediterranean Basin ecosystems Type Humid * Weeks Soil of drought <6 Alfi/Luvisols Vegetation beech-deciduous oak-pine Subhumid 6 -10 Nitosols evergreen oak - pine Semiarid 10 -25 Non-calcic browns desert grasslands Arid >25 Aridisols desert

Elev. (m) 1500 800 400 0 Elevation - vegetation relations (e. g. southern Italy) beech oak forest deciduous oak - pine forest sub-Mediterranean (=humid) evergreen oak - pine woodland semiarid grassland macchia Mediterranean (=subhumid) Mediterranean (=semiarid)

Montane pine forest, central Corsica

Evergreen oak forest, Mt Athos, N. Greece A monastic “templos” for over 1000 years

Vegetation - climate relations, California

Vegetation - topography relations, California N-facing S-facing

Common plants of the chaparral 1 5 2 1. Adenostoma fasciculatum 2. Heteromeles arbutifolia 3. Ceanothus verrucosus 4. Quercus dumosa 5. Cneroridium dumosum Dumos = ‘a bramble’ 3 4

Chaparral/maquis plants Common features: • microphylly (=small leaves) • sclerophylly (= hard “leathery” leaves with waxy surfaces) • aromatic foliage (herbs and spices) • woody stems • deep root systems

Plant seasonality rainfall drought

Water acquisition

Water conservation strategies

Sclerophyll leaf anatomy 1. Cutinized upper surface 2. Stomates in crypts 3. Tomentose openings to crypts 1 A B C A Ceanothus gloriosus [CA] B Banksia marginata [Aus. ] A Nerium oleander [Med. ] upper 2 3 lower

Plants of the coastal sage (CA) 2 1. Salvia apiana (white sage) 2. Salvia mellifera (black sage) 1 1

Plant tactics: allelopathy • Hypothesis that plants suppress competitors [and reduce herbivore attacks] by production of noxious chemicals ( e. g. aromatic foliage of Salvia; or oil-rich leaves of Eucalyptus which produce intense fires, kill competition) Bare zone and growth around Salvia patches in California inhibited grass Santa Inez Valley,

Post-disturbance succession or climax communities? evergreen oak - pine woodland Sclerophyllous woodland with broadleaved deciduous trees along streams. Trees <30 m maquis [Fr. ] matorral [Sp. , Ch. ] macchia [It. ] chaparral [Ca. ] garrigue [Fr. ] phrygana [Gr. ] coastal sage [Ca. ] mallee (Aus. ) Sclerophyllous shrubland: small trees <8 m high; often dense Increasing disturbance? Sclerophyllous shrubland: small shrubs <1 m high

Vegetation - climate relations, California

Community- environment interactions in shrublands

Thick litter buildup in mature chaparral

Wildfire in the chapparral

Burnt chaparral

Post-fire soil erosion e. g. sediment yield after 1989 Mt. Carmel [CA] forest fire

Fire and slope stability in southern California chaparral: links to the ENSO cycle El Niño (winter) La Niña (summer) normal (winter) r bic o h op r d e. g. 1997 -8 1998 -9 hy 1999* e lay s ow l df mu d n sa de sli 1999 -2000 * in the summer of 1999 x 2 average acreage burned in southern California

Post-fire recovery The majority of maquis and chaparral shrubs are capable of resprouting after fire (e. g. California lilac [Ceanothus tomentosus]). Fire reduces competition (by removing alleopathic litter) and opens up the canopy to allow germination of ‘fire annuals’.

Number of plants 5 m 2 Post-fire succession in chaparral* shrubs perennial ‘fire annuals’ *data from S. California; fire annuals are rare in Med. areas (e. g. Chile) with long fire recurrence intervals yrs

The Mediterranean as a “lost Eden”

Mediterranean vegetation: degradation to two endpoints? Oak woods maquis semi-phrygana semi-desert phrygana Based on Pantis, J. D. and Mardiris, T. A. 1992. Israel J. Bot. , 41, 233 -242 [F

maquis-phrygana mosaic (Crete)

Grazing intensity Goats at farmhouse near Antequera, Andalucia, Spain. (June 2007; photo: Dave Napthali)

Grazing and fire effects on Mediterranean vegetation Vegetation association (dominants) 1. 2. 3. 4. 5. Oak woodland (Quercus coccifera) Maquis (Q. coccifera-Thymus capitatus) Phrygana (T. capitatus - Ballota acetabulosa) Phrygana? (T. capitatus - Asphodelus aestivus) Geophyte semi-desert (A. aestivus ) Data: Pantis, J. D. and Mardiris, T. A. 1992. Israel J. Bot. , 41, 233 -242 [Table

Grazing and fire effects on Mediterranean vegetation - palatability Palatability according to local shepherds Data: Pantis, J. D. and Mardiris, T. A. 1992. Israel J. Bot. , 41, 233 -242 [Table 3]

Deforestation in the Middle East and Mediterranean world: the legacy of Gilgamesh The Epic of Gilgamesh, dating from about 2700 BC, celebrates the life of the legendary Babylonian king. On the 5 th clay tablet, Gilgamesh and his friend Enkidu (a former “wild-man”) travel for 7 days to confront the demon Humbaba, the guardian of the great Cedar Forest. After an epic battle, Humbaba is killed. Then: “they attacked the cedars - and while Gilgamesh felled the first of the trees of the forest Enkidu cleared their roots as far as the banks of the Euphrates”. After having felled the forest the heroes return on a cedar raft and use the tallest of the cedars to build a gate for the city of Uruk.

The legacy of Gilgamesh Deforestation resulted from: • Wood consumption for fuel (~90% of total? ) to supply domestic hearths, baths and industrial activities; shipbuilding, and military activities. Urban centres, metallurgical refineries and potteries placed a heavy demand on local fuel (charcoal) supply.

Deforestation in Cyprus Copper ores in Cyprus heavily exploited in Roman times, but mines functioned for about a millenium. Estimated that 5 -6 M tons of charcoal (from coppice) used to fuel smelting activities. Heavy SO 2 pollution may also have killed trees.

The legacy of Gilgamesh Deforestation resulted from: • Agricultural clearance - trees uprooted and burned down. Ashes used for fertilizer. • Pasturage - herdsmen cut branches to feed cattle and cut down trees to improve pasture for cattle and sheep. Pigs eat tree seeds, goats browse on young trees combined effect is permanent deforestation

Early anthropogenic deforestation in the eastern Mediterranean Jerf-el-Ahmar archaeological site N. Syria; ~11, 600 years old

9970± 100 12890± 160 olive evergreen oaks deciduous oaks cedar Deforestation in the Ghab valley, NW Syria pi es n Quat. Inter. 73/74, 127 -

Clearance at Sögöt, S. Turkey

Clearance of oak woodlands for pasture, subsistence agriculture and arboriculture above: subsistence farming (Andalucia) left: Olives (Jaen) Photos: Dave Napthali, June 2007

Initiation of land degradation in the Mediterranean basin Goudie, A. 1992. Environmental Change. Oxford

The effects of deforestation On the landscape of Attica (central Greece), Plato commented: “what now remains compared with what then existed is like the skeleton of a sick man, all the fat and soft earth having wasted away, and only the bare framework of the land being left” Critias 111 B

South Holocene valley fills in Greece North Are these episodes of erosion associated with phases of climate change, or are they exclusively

Soil erosion and valley filling: e. g. Ephesus, W. Turkey

A “heretical” viewpoint: Grove and Rackham “some of their interpretations are "heretical" and not shared by many of their. . research colleagues. … These include the notion that there is little if any desertification going on in (Mediterranean) Europe …; that Mediterranean Europe's badlands and most of its erosion is natural and not the result of human action (here they endorse Vita-Finzi's ideas); that the human shaping of the landscape of Mediterranean Europe was for the most part complete by the Bronze Age, and that human agency has not done much to change it since-until perhaps the last thirty years when bulldozers have been let loose. ” Abstracted from the review by J. R. Mc. Neill, 2001. Environmental

Deserted village and invading pines, western Crete (photo: Oliver Rackham, 1989) Rural depopulation, mountains of Aragon, Spain. (data: Collantes, F. , Pinilla, V. 2005. Rural History, 15, 149 -166. ) "Big fires are a predictable result of rural depopulation, land abandonment, increase of wild vegetation, modern forestry, legislation against fire, and the growth of. Fire in the European Mediterranean. a fire-fighting Rackham, O. 2003. Aridlands Newsletter, No. 54, November/December industry. "