Contribution to the study of the Greek flora: Flora and vegetation of the Enousses (Oinousses) islands (E. Aegean area)

FLORA (1994) 189 69 -78

© by Gustav Fischer Verlag lena

Contribution to the study of the Greek flora: Flora and vegetation of the Enousses (Oinousses) islands (E. Aegean area) M. PANITsa, P. DIMOPOULOS, G. IATRou, D. TZANOUDAKIS* University of Patras, Department of Biology, Division of Plant Biology, 265 00 Patras, Greece Accepted: September 28, 1993

* To whom all correspondences should be sent

Summary Enousses is an islets group situated eastwards of Chi os island (E. Aegean, Greece), which remained quite unexplored tloristically. Based on our herborizations, 270 taxa belonging to 51 families and 204 genera of spermatophytes are listed for the tlora of this area. The tlora of the area is analysed and the plant species recognized are classified into 9 chorological groups and 5 life form types. The results of this analysis confirm the Mediterranean character of the tlora since the Mediterranean elements and the therophytes are predominant in the area. The vegetation of all islets is characterized by the dominance of phryganic communities, and the three "vegetation zones" distinguished (littoral, sublittoral and interior ones) are described. Geological history, geography, microecological differences, random events and human activities seem to be the most important factors related to the diversity of the tlora and vegetation observed in our area. Special attention has been given to human interference, since in the bigger islets of the group -which are used for agriculture, grazing etc. - high percentages of Therophytes and Leguminous species have been observed (indexes of disturbance). On the contrary, in the smaller ones, which are not affected by human activities (i.e. Vatopoula), these indexes show deviate values. For this reason, the importance of the small islets ecosystems in tloristic and ecological studies in the Aegean area is pointed out. Key words: Flora, vegetation, Greece, Aegean area, islets ecosystems

1. Introduction In the framework of a research project dealing with floristicand phytogeographical studies in the Aegean area, emphasis has been given to the small and isolated islets of the eastern Aegean, since many of them remained quite unexplored, from a floristic point of view. The area under study belongs to this category, since there is no floristic or any other related information concerning the Enousses island group. There are no references neither in the "Flora Aegaea" (RECHINGER 1943) nor in the "Flora of Turkey" (DAVIS 1965-1985). Moreover, no information is included in MEIKLE'S survey on the "Flora of Chi os" (MEIKLE 1954).

Enousses is an eastern Aegean Greek islet group lying in the sea channel which separates the NE coast of Chios Island from the western coast of Asia minor (38 20 o N, 26 08°E). The nearest coast of Chi os lies at a distance of 2 km to the west, while that of Asia Minor ca. 8 km to the east of the island group. The island group consists of 6 islets namely: Enoussa (E) (Oinoussa), Panaghia (Pa), Vatos (V), Vatopoula (VI), Pontikos (Po) and Archontonisso (A). The main islet is Enoussa which covers an area of ca. 14 km 2 and has a maximum length of ca. 8 km. The others are much smaller covering a surface of ca. 3 km 2 with maximum lengths varying from 3 km to 100 m (Fig. I, Table 3). FLORA (1994) 189

69

F'

151. ENOUSSES

38"30'

o

2

,

6

4

Fig. I. Geographical position of the Enousses islets group, in Greek area and detailed map of the area studied. I: Enoussa (E), 2: Panaghia (Pa), 3: Vatos (V), 4: Pontikos (Po), 5: Vatopoula (Ve), 6: Archontoniso (A)

Table I: Life form spectrum of the total nora of the Enousses islets.

Life forms

taxa

%

Chamaephytes Ch. fruticose Ch. suffruticose Ch. reptant

(Ch) [Chfrut] [Chsuffr] [Chrept]

18 6 11

6.7

Geophytes G. rhizomatose G. bulbose G. parasite

(G) [Grhiz] [Gbulb] [Gpar]

31 8 22

11.5

Hemicryptophytes H. scapose H. caespitose H. scandent H. rosulate

(H) [Hscap] [Hcaesp] [Hscand] [Hros]

35 16 11

13.0

Phanerophytes Nano-ph. Nano-scandent

(Ph) [Nph] [Nscand]

17 16

6.3

Therophytes Th. scapose Th. caespitose Th. rosulate Th. reptant Th. parasite

(Th) [Thscap] [Thcaesp] [Thros] [Thrept] [Thpar]

169 157 2 4 3 3

62.5

270

100

Total

70

FLORA (1994) 189

7

Geologically, the whole islet group is more or less uniform since it consists of low-grade metamorphic rocks. Schists of pelitic nature predominate in the greatest part ofthe main islet, while in the smaller islets and the adjacent SE part of Enoussa the psammitic rocky formations are predominant. Conglomerate layers, marls and marly limestones are also represented but with a minor role in the geological structure of the area. Regarding the topography of the area, each islet consists of one, rarely more, low altitude land formation, with the highest point reaching 182 m above sea level on the Enoussa islet. The hill slopes are more or less of gentle gradient and as a consequence, deep valleys, gorges, vertical rocky systems and other similar specialized biotopes have not been observed. Sandy beaches are also rare in the area and these are significantly limited when present. From the six islets studied, only the biggest one, Enoussa, has been inhabited since ancient times. Based on historical reports and on the islets name as well r"oivos" (oinos) means wine in Greek] it is concluded that vineyards were the predominating agricultural activity. Today the cultivations are very restricted and only some small patches exist in the western part of the islet, especially around the monastery of Panaghia. Contrary to viticulture, stock farming persists nowadays and intense grazing takes place on the main islet and the smaller ones as well. Actually one of them, Archontonisso, has been transformed into a sheepfold, and as a result, only a few perennial shrubs are still surviving on it. The bioclimate ofthe area is illustrated by the coefficient of EM BERGER (EMBERGER 1955, MAVROMMATIS 1980) and the ombrothermic diagrams of BAGNOULs-GAussEN (1957). The

climatic data come from the meteorological station of the neighbouring island of Chios (altitude 3 m, period 28 years: 1932-1941 & 1951-1971). According to the EM BERGER'S coefficient, the area belongs to the sub-humid bioclimatic stage with mild winters, almost free offrost and snow, (Fig. 2 & 3). According to BAGNOULS & GAUSSEN (I.c.) the dry period lasts almost seven months (from mid April to late October).

n::

Pmm 180

90

160

80

200

150

HU MID

3

o

J

F

M

A

M

J

J

A

s

o

N

o

f'ig. 2. Ombrothermic diagram of Chios meteorological station.

Mild

winter

7

10

W.rm

m

winter

Fig. 3. E\1BERGERS pluviothermic c1imagramme for Enousses islets (according to climatic data of Chios meteorological station).

2. Materials and methods The plant list given below is based essentially on our own collections since no other tloristic information was available. Our team visited the area for the first time in April 1989 from 12th to 15th; members of the team made repeated collections in early May and in October of 1990. Vouchers are deposited at UPA. The sequence of families. genera and species in the plant list is alphabetical. The nomenclature is in accordance with DAVIS ( 1965-1985) and. where possible. with GRITTER et al. ( 1984-1989). These basic tloras have also been used as main sources regarding the geographical distribution of the taxa. For the chorological analysis of the tlora the following nine chorological types have been recognized. I. Cosmopolitan. 2. Subcosmopolitan. 3. Paleotemperate.4. MediterraneanTuranian. 5. Mediterranean-Atlantic. 6. Euri-Mediterranean. 7. StenoMediterranean. 8. Mediterranean-Irano-Anatolian. 9. East-Mediterranean.

The chorological types 1-7 are in accordance with PIG'ATTI ( 1982). For the last two chorological types (8 & 9) the corresponding definitions are as follows: Med-Irano-Anat.~ In this chorological type are included tloristic elements (taxa) with their distribution center in W. and C. Asia but their distribution range expanded westwards to the Mediterranean area (cL C\RLSTRo\l 1989). E. Med.: In this chorological type are included taxa with their distribution range restricted to the coasts of the East Mediterranean i.e. extending from eastern Italy to Lebanon (cL DAVIS 1965). With regard to the ranking of taxa into life-form categorics and the production ofa life-form spectrum, R.,\I"KIAERS system (RAI'WIo\FR 193-1) was followed.

3. Results 3.1. Flora As a result of our field work, 270 taxa distributed to 51 families and 204 genera are recorded from the Enousses island group, for the first time. In the species list given below are not included the perennial species Pinus brutia, Nerium oleander, Nicotiana glauca, Ricinus communis and Eucalyptus .Ip., whose presence has

been noticed around the harbour-village of the main islet Enoussa. These taxa have not been taken into account in the floristic analysis (Tables I - 4) since, obviously, these are not native in the localities where we observed them. From the standpoint of richness of species, Legumil1osae, Compositae and Gramineae are represented by 50, 37 and 30 taxa respectively. These three families comprise 43 % of the taxa found in the studied area. FLORA (1994) 189

71

-

For the chorological analysis we have taken into consideration 264 taxa which are properly determined. By means of this analysis, nine chorological groups have been distinguished (Table 4) and lead to the following ascertainments: - The East Mediterranean group includes the mostly narrow distributed element of the examined chorological spectrum and represents 21,2 % (56 taxa) of the total flora. - The total number of Mediterranean elements (E. Med., St. Med., Eu. Med.) participating in the Enousses flora is 183 (68,1%). -The other chorological groups which contain more widely distributed taxa, are represented by lower percentages. From a floristic point of view, the majority of the species recorded from the Enousses islets have already been reported either from the neighbouring islands of Chios (RECHINGER 1943, MEIKLE 1954, DAVIS 1965-1985) or from some other eastern Aegean islands (DAVIS i.c., CHRISTODOULAKIS 1986). Regarding the life-form spectrum of the Enousses flora, from Table I it appears that the percentage of the Therophytes (62,5 %) is much higher than of all the other life-forms. This is in accordance with the character of the Mediterranean bioclimate prevailing in the studied area (Intense Thermomediterranean) and the long dry period (MAVROMMATIS 1980).

taxa

Ch

50

2.0 10.8

Gramineae

37 30

Labiatae Caryophyllaceae Liliaceae

12 12 12

50.0

Cruciferae Umbelliferae Plantaginaceae

10

G

Life-forms (%) H Ph

Number of taxa

Percentage

Eastmediterranean (E. Med.) Stenomediterranean (St. Med.)

56

21.2

38

Eurimediterranean (Eu. Med.) Mediterranean-Atlantic (Med.-Atl.)

89

14.4 33.7

10

3.8

3

l.l

Chorological group

(Med.-Turan.) Mediterranean-Irano-Anatolian (Med.-Ir. Anat.) Paleotemperate (Paleotemp.) Subcosmopolitan (Subcosmop.) Cosmopolitan (Cosmop.)

24

9.1

36

13.6

I 7

0.4 2.7

264

100

3.2. Plant list

Th Gymllosperl11ae

Leguminosae Compositae

8 6 6

Scrophulariaceae

4.0 2.7 6.7

4.0

21.6 23.3 25.0 16.7

90.0 64.9 70.0

8.3

16.7 83.3

100.0 12.5

12.5

10.0 12.5

90.0 62.5

16.7

83.3 66.7

33.3

Ephedraceae Ephedra call1py/opoda c.A. MEYER: NPh, E. Med.; VI Allgiosperl11ae Dicoty/edolle.l· Allacardiaceae Pisracia/ellti.I'('lIS L.: NPh, St. Med.; A, E, Pa, Po, V. VI P. terehillthlls L. subsp. pa/aestilla (80155.) ENGI.ER: NPh, E. Med.; E Arist%chiaceae Arist%chia/JlII1'if;,/ia SM.: Gbulb, E. Med.; E. Pa, V Boragillaceae Echiwll p/alltagillelll11 L.: Thscap, Eu. Med.; E, Pa, V

Table 3. Representation of the Leguminosae (Leg)and Therophytes (Th) in the tloras oftive islets of the Enousses group in comparison with the total tlora of the area. The islets are arranged according to their decreasing size and for abbreviation see tigure I. Islet

Surface

Taxa

Leg

Th

~i-

%

172 125

16.3 25.6

61.3

ca. 0.3 km 1 ca. 0.04 km 1

18 83 50

11.1 10.7 12.2

km 1

270

18.5

E

ca. 14.7 km 1

Pa Po V

ca. 2.7 km 1 ca. 0.4 km 1

VI Total

72

ca. 18.14

FLORA (1994) 189

77.9 38.9 52.5 36.0 62.5

%

Mediterranean-Irano-Turanian

Abbreviations used: i) Collections sites (islet): A=Archontonisso, E=Enoussa. Pa=Panaghia, Po=Pontikos, V=Vatos, VI=Vatopoula. ii) Life forms: See table I iii) Chorology: See table 4.

Table 2. The ten richest in taxa families of the Enousses tlora.

Families

Table 4. Chorological spectrum of the Enousses tlora.

Myosotis ralllOSissil11a ROCHEL subsp. ral11osissil11a: Thscap, Paleotemp,; V Call1pallll/aceae Call1p(lllll/a erillus L.: Thscap, Eu. Med.; E Legol/sia pelltagollia (L.) DRUCE: Thscap, E. Med; E, Pa Cappal'llceae Cappari.l· spillosa L.: NPh, Eu. Med. (Atl.); E Caryophy/lt/('eae Arellaria/eproclados (REICHE;>;B.) GU55.: Thscap. Paleotemp.; Pa CercWilll11 COl11atlll11 DESV.: Thscap, E. Med.; Pa C. g/olllel'llflll11 THUII.\..: Thscap, Paleotemp.; E. Pa Parollycflial11acrosepa/a 80155.: Hcaesp, E. Med.; E, Po, V, VI

--,

Petror/wgia velutina (Guss.) P.W. BALL. & HEYWOOD: Thscap, Eu. Med .. ; E,Pa Polrcarpon tetraphyl/ulIl (L.) L.: Thscap, Eu. Med.; E, Pa, V Silene colorata POIRET: Thscap, Eu. Med.; E, Pa S·ftlharia (L.) SM.: Hscap, E. Med.; VI S. gallica L.: Thscap, Paleotemp.; Pa S. sedoides POIRET: Thscap, SI. Med.; E Spergularia hocconei (SCHEELE) GRAEBNER: Thscap, Eu. Med. (Atl.); E, Pa Stel/aria pal/ida (DCMORT.) PIRE: Thscap, Paleotemp.; E, V Chenopodiaceae Atriplex halimus L.: NPh., Med.-Atl.; V, VI A. prostrata DC.: Thscap, Paleotemp.; Po Chenopodiulll murale L.: Thscap, Cosmop.; V Salsola kali L.: Thscap, Paleotemp.; Po Cistaceae Cistus creticus L.: NPh, E. Med.; E, Pa, Po, V, VI C. salvii/illius L.: NPh, Eu. Med. (lr. Anal.); A, E, VI Fumana arabica (L.) SPACH: Chsuffr, Eu. Med. (lr. Anat.); E, V Tuberaria guttata (L.) FOCRR.: Thscap, Med.-Atl.; E, Pa Compositae Aetheorhi:a bulhosa (L.) CASSo suhsp. microsepala RECH.: Gbulb, E. Med.; E Andrmla integri/(Ilia L.: Thscap, SI. Med.; E Anthemis chia L.: Thscap, E. Med.; E, Pa A. rigida BOIss.: Thscap, E. Med.; E, Pa, V, VI Asteriscus aquaticus (L.) LESS.: Thscap, St. Med.; E Atractylis cancel/ata L.: Thscap, Eu. Med.; E A. gllmmi(era L.: Hros, SI. Med.; V Bel/ium minlltum L.: Thscap, E. Med.; E, Pa Calendula arvensis L.: Thscap, Med.-Ir. Anat.; Pa, V Carduus /'-"c/lOcephallis L.: Thscap, Med.-Ir. Anal.; E, Pa, V Centaurea spinosa L.: Chfrut, E. Med.; E, Pa, Po, V, VI Cllrrsanthel1lul1l cOr!lnariul1l L.: Thscap, Eu. Med.; E Crepis dioscoridis L.: Thscap, E. Med.; V C. commutata (SPRENG.) GREUTER: Thscap, E. Med.; E, Pa C. multiflora SM.: Thscap, E. Med.; E, Pa Crupina crupinastrulll (MORIS) VIS.: Thscap, Eu. Med.; E Dittrichia visco,l'a (L.) GREUTER: Hscap, Eu. Med.; E Echinops viscosus DC. subsp. visCIIsus: Hscap, E. Med.; V Filago aegaea WAGENITZ subsp. aristata W AGl'NITZ: Thscap, E. Med.; E F. eriocephala Guss.: Thscap, Eu. Med.; Pa Hedypnois cretica (L.) DUM-COCRSET.: Thscap, Eu. med.; E, Pa, V Helichn'sum stoeciws (L.) MOENCH subsp. barrelieri (TEN.) NYMAN: Chsuffr, E. Med; Pa, Po, VI H. orientale (L.) DC.: Chsuffr, E. Med.; V HypocilOeris achyrophorus L.: Thscap, St. Med.; A, E, Pa, V, VI H. glabra L.: Thscap, Paleotemp.; E, Pa, V Leontodon tuberosus L.: Hros, SI. Med.; E, Pa, VI Log.fia gal/ica (L.) COSSON & GERM.: Thscap, Eu. Med.; Pa, V, P/wgnalon graecul1l BOlss.: Chsuffr, E. Med.; Po, V, VI Pici1l1omon acarna (L.) CASS.: Thscap, Eu. Med.; E Reichardia picroides (L.) Roth: Hscap, St. Med.; E, V Senecio I'ulgaris L.: Thscap, Paleotemp.; Pa, VI Sonchus oleraceus L.: Th (H) scap, Paleotemp.; V Taraxacum aleppicum DAHI.ST.: Hros, E. Med.; E Taraxacum sp. (secl. Scariosa) ed.: SI. Med.; E Tragopogon porrijillius L.: H (Th) scap, Eu. Med.; E, V Tolpis barbata (L.) GAERTNER: Thscap, SI. Med.; E, Pa Urospermlllll picroides (L.) SeDP.: Thscap, Eu. Med.; E, Pa, V

COIII'oll'ula{'ecle Conl'Oit'ulus elegantissimus MILLER: Hscap, SI. Med.; A, E Cuscllta sp.: Thpar; E Crassulaceae Sedum pallidum MB.: Thscap, E. Med.; E S. rubens L.: Thscap, Eu. Med.; E Umbilicus /lOrbll1talis (Guss.) DC.: Gbulb, Eu. Med.; E, Pa, V U. rupestris (SALlSB.) DANDY: Gbulb, Med.-Atl.; Pa Cruci(erae Biscutel/a didwlla L.: Thscap, Eu. Med.; E, VI Cakile maritima ScoP.: Thscap, Paleotemp.; E Calepina irregularis (Asso) THELL.: Thscap, Med.-Ir. Anal.; Pa Capsel/a bursa-pastoris (L.) MEDlK.: Thscap, Cosmop.; Pa Malcolmia chia (L.) DC.: Thscap, E. Med.; VI M . .f7exuosa (SM.) SM.: Thscap, E. Med.; E, Pa, V Matthiola tricuspid{{fa (L.) R. BR.: Thscap, SI. Med.; E Sisymbrium ()flidnale (L.) Seop.: Thscap, Paleotemp.; E S. orientale L.: Thscap, Med.-Ir. Anal.; E Teesdalia coro)lOPi(olia (J.P. BERGERET) THELL.: Thscap, Eu. Med.; E Elaeagnaceae Elaeagnus angusti/(Ilia L.: NPh, E. Med.; V Ericllceae Erica nUlIlipuliflora SALlSB.: NPh, E. Med.; E

Euphorbiaceae Euphorhia helioscopia L.: Thscap, Cosmop.; E E. peplis L.: Thrept, Eu. Med.; E. Mercurialis ({)1I1ua L.: Thscap, Paleotemp.; V Frankeniaceae Frankenia hirsuta L.: Chsuffr, Med.-Ir. Anal.; VI Gelltian(lceae Blackstonia peifoliata (L.) HUDSON subsp.l'e/:/()liata: Thscap, Eu. Med.; V

Geraniaceae Erodium cicutariul1l (L.) L'HER.: Thscap, Paleotemp.; E, V E. mosciwtum (L.) L'HER.: Thscap, Eu. Med.; Pa Geraniul1l dissectum L.: Thscap, Paleotemp.; Pa, V G. mol/e L. subsp. l1lol/e: Thscap, Paleotemp.; E Gutti(erae Hypericum pe/.foratum L.: Hscap, Paleotemp.; E, V Labiatae Bal/ota acetabulosa (L.) BENTHAM: Chfrut, E. Med.; E, Pa CoridothYl1lus capitatus (L.) REICIIENB. til.: Chfrut, SI. Med.; E Lalllium amplexicaule L.: Thscap, Paleotemp.; E, Pa Lavalldula stoechas L.: NPh, SI. Med.; E, Pa, V Marrubium I'ulgare L.: Hscap, Paleotemp.; E Origanul1l onites L.: Chsuffr, E. Med.; E Prasium mq;us L.: Chfrut, SI. Med.; Pa, VI Sait'ill I'erbenaca L.: Hscap, Med.-Atl.; V Safllreja juli({)w L.: Chsuffr, SI. Med.; E S. thYl1lbra L.: Chfrut, E. Med.; E Sideritis cU/Tidens STAPF: Thscap, E. Med.; Pa Stachys cretica L. subsp. smyrnaea RECH. fil.: Hscap, E. Med.; E

FLORA (1994) 189

73

--,..

Legulllillosae (Fabaceae) Allthyllis hermalllliae L.: Chfrllt, SI. Med.; E, Pa Astragalus halllosus L.: Thscap, Ell. Med.; Pa Astragalus pelecil1l1s (L.) BAR:-.JEBY: Thscap, Ell. Med.; E, Pa Bitumillaria bitumillosa (L.) STlRTO:-:: Thscap, Ell. Med.; Po, V Calicotome I'il/osa (POIRET) LI:-.JK: NPh, SI. Med.; E, VI Hymellocarpos circillatlls (L.) SAYL: Thscap, Ell. Med.; E, Pa Lathyrus cicem L.: Thscap, Med.-Ir. Anat.; E L. setifillills L.: Thscap, Ell. Med.; E L. sp/wericus RETZ.: Thscap, Paleotemp.; Pa Lells nigri(,([lls (M.B.) GODRON: Thscap, Ell. Med.; E Lotus coimbricellsis BROT.: Thscap, SI. Med.; Pa L. aeticus L.: Thscap, SI. Med.; Pa, VI L. edlliis L.: Thscap, SI. Med.; E, Pa, V L. omit/wpodioides L.: Thscap, Ell. Med.; E L. cf. peregrilllls L.: Thscap, E. Med.; Po L. tetragllllOlolms L.: Thscap, Ell. Med.; V Lupillus {//1gustijilliliS L.: Thscap, SI. Med.; Pa L. lIIicrallthus GLSS.: Thscap, SI. Med.; E, V, VI Medi('([go corollaf(/ (L.) BARTAL.: Thscap, Ell. Med.; Pa M. discifimnis DC.: Thscap. SI. Med.; E, Pa M. lIIollspeliaca (L.) TRALTY.; Thscap, Ell. Med.; Pa M. orhicularis (L.) BARTAL.: Thscap, Med.-Ir. Anal.; E, Pa, V M. praecox DC.: Thscap, Ell. Med.; E, Pa M. tuberclilata (RETZ.) WII.LD.: Thscap, SI. Med.; Pa Ollobrl'chis mput-gal/i (L.) LAM.: Thscap. Ell. Med.; E, Pa Ollollis pllsil/a L.: Hscap, Eu. Med.; Pa Omithopus compresslls L.: Thscap, SI. Med.; E. Pa O. pill/wtus (MILLER) DRLCE: Thscap, Med.-Atl.; Pa Scorpiurus IIIl1ricatus L.: Thscap, Ell. Med.; Pa, VI SpartiulII jUllcell1ll L.: NPh. Ell. Med.; E. VI Trijilliwll allgustijillilllll L.: Thscap. Ell. Med.; Pa T. {//'I'ellse L.: Thscap, Paleotemp.; Pa T. bul/atum BOIss. & HALSSK~.: Thscap, E. Med.; E T. campestre SCIIREBIX Thscap. Paleotemp.; A. Pa, V T. gralldij/orlllll SCIIREBER: Thscap, E. Med.; E T. hirflllll ALL.: Thscap, Ell. Med.; E, Pa T. Iligrescells VIY.: Thscap, Eu. Med.; E, Pa T. pillilare BOlss.: Thscap, E. Med.; E T. resupillatulII L.: Thscap, Med.-Ir. Anal.; E T. s('([brulll L.: Thscap, Ell. Med.; V, VI T. SPIlIIIOSIIIIl L.: Thscap. Eu. Med.; Pa T. stel/afl/lI1 L.: Thscap. Eu. Med.; E, Pa T. sllbterralleum L.: Thscap. Ell. Med.; E T. ullij/orulll L.: Hcaesp. SI. Med.; E, Pa, V Trigonel/a balall.we BOlss. & RITTER: Thscap. E. Med.; Pa Vicia hybrida L.: Thscap, Ell. Med.; E V. lathrroides L.: Thscap, Ell. Med.; Pa V. sati\'{/ L. sllbsp. Iligra (L.) EIIRII.: Thscap. Cosmop.; E. Pa. V V. \·il/osa ROTH subsp. erioc1ll7)(1 (HACSSK:-J.) P.W. BALL: Thscap. Ell. Med.; E sllbsp. lIIicrophyl/a (DUM.-URYILLE) P.W. BALL: Thscap, ElI.-Med.; Pa Lil1(fCeae

Lilllllll trigYllll1ll L.: Thscap, Ell. Med.; E. Pa Mall'aceae Mal\'{/ cretica CAY.: Thscap. SI. Med.; Pa M. pan'ij/ol'(l L.: Thscap, Ell. Med.; E Mal\'{/ sp.: Thscap; E MOl'(lceae Ficus ('([rica L.: NPh, Med.-Ir. Anal.; E

74

FLORA (1994) 189

Myrtacea(!

Mwtus

COllII/HlIlis

L.: NPh, Med.-Ir. Anal.; E

Oleaceae Olea europaea L. sllbsp. oleaster (HOFFMA:-;~S. & LINK) NEGODI: NPh, Ell. Med.; A. E, VI Oroballc/wceae Orohallche plibescells DLM.-URYILLE: Thpar. Ell. Med.; E Oroballche cf. reticulata WALLR.: Thpar, Eu. Med.; E, V Oxalidaceae Oralis pes-caprae L.: Gblllb, Cosmop.; E. Pa, V Pal}(l\'eruc(!a(!

FlIIl1aria dellsif/ora DC.: Thscap, SlIbcosmop.; Pa F. judaica BOlss.: Thscap, E. Med.; Pa F. lIlacro('([pm PARL.: Thscap. E. Med.; E. Pa Pap{/\'er argemolle L.: Thscap. Med.-Ir. Turan.; E, Pa P. duhium L.: Thscap, Paleotemp.; E Plli/ltligi !l{lC'Cae

Plalltago {(ji-a L.: Thscap. Med.-Ir. Anal.; E, Pa P. bel/ardii ALL.: Thros, Ell. Med.; VI P. cretiC{{ L.: Thros, E. Med.; E P. lagopus L.: Thros, Eu. Med.; E, Pa P. lallceolata L.: Hros, Paleotemp.; E. VI P. lI'eldellii RElCIIE~B.: Thros, Med.-Ir. Anal.; E, Pa, VI Plumbagilwceae Arllleria cariellsis BOlss.: Hros. E. Med.: Po, VI LiIllOlliulII glllelillii (WILI.D.) O. KL:\TZE: Chsllffr. St. Med.: VI L. graecum (POIRET) RECII. fil.: Chsllffr. E. Med.; A. E. Po. VI L. sillUafl/lII (L.) MII.I.ER: Hscap. Ell. Med.: Po. V L. \'irg{/fulII (WILLD.) FOl'RR.: Chsuffr, Med.-Atl.; E Pol.\'g()lll/('C{Il!

Rumex buccphalophol'lls L.: subsp. buccphalophol'lls: Thscap. Ell. Med.: Pa. V subsp. gal/icus (STEINH.) RECH. fil.: Thscap. Ell. Med.: E. Pa. V R. pulcher L. subsp. \\'(}odsii (DE NOT.) ARCANGu.l: Hscap. Ell. Med.: E R. fl/bcrosus L. sllbsp. tuherosus: Gblllb. Med.-Ir. Tliran. E. Po. V. VI Primllllu'cae

Allagal/is (/I'I'ellsis L.: Thrept. Paleotemp.; Pa. V Asterolillolllilllllll-stel/afl/l11 (L.) DLHY: Thscap. Eu. Med.: E. Pa. V Rafj/csia('cac Cytilllls hypocistis (L.) L. sllbsp. cillsii NYMA:-J: Gpar, Ell. Med.: E Rall1ll1culacme Clematis cirrhosa L.: Nsc,\),Il. St. Med. (Atl.): E RlI1lllllCllllls lII11rgil111tlls Dl'\1.-URYII.I.E: Thscap. E. Med. (lr. Anat.): Pa Rosliceae

Sarcoporerilllll spil10sulII (L.) SPAU!.: NPh. E. Med.: A. E. V. VI RlIiJillcelle Catilllll aparil1c L.: Thscap. Paleotemp.; E, Pa C. lIIurale (L.) ALL.: Thscap, Eu. Med.: Pa C. recur\'1llll REG.: Thscap. E. Med.: E. Pa

--,..----

Sherardia arvcllsis L.: Thscap, Paleotemp.; E, V Va/allfia mura/is L.: Thscap, SI. Med.; Pa, V Sal1ta/accac Thesiulll bergeri Zece.: Chrept, E. Med.; E Scrophu/ariaccae Bel/ardia trixaRo (L.) ALL.: Thscap, ELI. Med.; V Lil1aria peiis.lcrial1a (L.) MIl.l.ER: Thscap, Med.-Atl.; E, Pa Misopates orollfiul11 (L.) RAmI.: Thscap, ELI. Med.; Pa Scrophu/aria cal1il1a L. sLlbsp. bic%r (SM.) GREUTER: Hscap, E. Med.; E Verbascum sp.: Hscap; E Verollica cnllba/aria BODARD: Thscap, ELI. Med.; Pa So/allaceac Mal1draRora autwlllla/is BERTOL.: Hros, ELI. Med.; V Ulllbel/i/erae Crithmum maritimulII L.: ChsLltlr, Med.-Atl.; V, VI Daucus il1l'O/ucratus SM.: Thscap, E. Med.; Pa Er\'llgiulllmaritilllum L.: Grhiz, Med.-Atl.; VI Fcm/a commullis L. sLlbsp. g/auca (L.) Rouy & CAMes: Hscap, ELI. Med.; E Lagoccia CUlllilloides L.: Thscap, ELI. Med.; E Sc{//u/ix pectell-l'elleris L.: Thscap, Paleotemp.; E Tord,l'/iulII apu/ulll L.: Thscap, SI. Med.; E, Pa, V Tori/is /cptopilyl/a (L.) REICHB.: Thscap, Med.-Ir. Anal.; Pa UrficlIc(!a(!

Parietaria cretica L.: Thrept (Hscap), E. Med.; Pa

Valerianaceae Va/eriallel/a discoidca (L.) LOIs.: Thscap, ELI. Med.; E, Pa V. dClltata (L.) POLL.: Thscap, Med.-Atl.; E Verbcllaceac Vitex agllus-('(/stus L.: NPh, Eu. Med.; E M,J/1o('oty/edolles AlIlary//idaceac Narcissus scmtillll.l· L.: GbLllb, SI. Med.; V Anu'(!a(!

Ari.mrulIIl'II/garc TARG.-Tozz.: Grhiz, ELI. Med.; VI

C."per(/c(!ae Carex sp.: Hcaesp; Pa, V Sci/pus h%schoellus L.: Grhiz, Paleotemp.; V

B. tectorulI1 L.: Thscap, Paleotemp.; E, Pa Catapodiull1 riRidulIl (L.) C.E. HUBBARD: Thscap, ELI. Med.; Pa, VI Dactl'/is R/oll1erata L.: Hcaesp, ELI. Med.; A, E, Po, V, VI E/Yll1us/clrctus (VIV.) RL'~E\IARK (s.I.): Grhiz, ELI. Med.; VI HordeulIl murillulIl L.: Thscap, Med.-Ir. Anal.; E, Pa Hyparrhcllia hirta (L.) STAPF: Hcaesp, Med.-Ir. Anat.; E Lagurus omtus L.: Thscap, ELI. Med.; E, Pa, Po, VI Lo/ium rigidum GACDIN: Thscap, Med.-Ir. Anal.; E, Pa Me/ica milluta L.: Hcaesp, St. Med.; VI Parapho/is illcun'a (L.) C.E. HeBBARD: Thscap, Med.-Ir. Anal.; Pa, VI Phragmites australis (CAV.) TRI~.: Grhiz, Cosmop.; V Poa ill/irma KC~TH: Thscap, Med.-Ir. Anal.; E, VI P. tril'ia/is L.: Hcaesp, Paleotemp.; V P. bulbosa L.: Hcaesp, Paleotemp.; E Rostaria cristata (L.) TZVEl.EV: Thcaesp, Med.-Ir. Anal. Pa, V Stipa capellsis THU!'B.: Hcaesp, Med.-Ir. Anal.; Pa TrachYllia distachya (L.) LI~K: Thscap, Med.-Ir. Anal.; E, Pa, V Vu/pia myuros (L.) c.c. GMELlN: Thcaesp, Paleotemp.; Pa Iridaceae Crocus sp.: GbLllb; E JUllcaceae JUIICIIS hc/drcichiallus MARSSON

sLlbsp. he/drcichial1l1s: Hcaesp, E. Med.; Pa, V Li/iaccae Al/iulIl cupalli

RAFI~.

sLlbsp. ilirtomgilllltulII

(KL:~TH)

STEAR:-J: GbLllb, ELI. Med.; E, VI

A. .t1{/\'1II1l L.

subsp. tauriculII (BESSER ex REICHB.) STEAR!': GbLllb, St. Med.; E A. guttall/Ill STEVE:-; sLlbsp . .mrdoulII (MORIS) STEAR~: GbLllb, SI. Med.; E, V A. lIeapo/itallulII CVR.: Gbulb, SI. Med.; E A. roSCUIll L.: Gbulb. SI. Med.; E. V Asparagus acutijil/ius L.: Grhiz, SI. Med.; VI Asphode/us aestil'lls BROT.: Grhiz, ELI. Med.; E. Po, V, VI Cagea gracca (L.) A. TERRACe.: GbLllb, E. Med.; Pa, V Muscari COIIIOSIIIII (L.) MILLER: GbLllb, ELI. Med.; E M. ll'cissii FREY~: GbLllb, E. Med.; E, Pa, Po, V, VI Omithoga/ulll cf. arlllelliaculll BAKER: GbLllb, E. Med.; E Urgillea lIlaritima (L.) BAKER: GbLllb, ELI. Med.; E Orchidaccac Lilllodol'lllll aborti\'lllll (L.) SWARTZ: Grhiz, ELI. Med.; E NeotillCll lIIacu/ata (DESF.) STEAR~: Gbulb, ELI. Med.; V Ophrl's ,IIleIfleri Fl.EISCHM.: GbLllb, E. Med.; V Orchis S{//Icta L.: GbLllb, E. Med.; E SCl'llpia.l· I'IIIIlCl'llcca (BeRM. fil) BRIG. sLlbsp. /ax(flol'll (S()(i) GOl.z. & REINHARD: GbLllb, E. Med.; E. Pa, V

Gramilll!lIl:!

I

I

I

I j I

'1

Acgi/ops Cl'/illdrica HOST: Thscap, Med.-Ir. TLlran.; E Aim e/egallfissi111a SCHUR: Thscap, ELI. Med.; E, Pa A11Ihoxa11lhulll odoratulII L.: Hcaesp, Paleotemp.; E Al'ella barbata Pon: Thscap, Med.-Ir. Anal.; E, Pa, VI A. stcri/i.l· L.: Thscap, Med.-Ir. Anal.; E Bri~a lIlaxillla L.: Thscap, ELI. Med.; E, Pa, V, VI B. millor L.: Thscap, Cosmop.; V Brolllus a/opecuros POI RET: Thscap, ELI. Med.; Pa B. illterllledills Guss.: Thscap, ELI. Med.; E, Pa, V B. lIIadritellsis L.: Thscap, Paleotemp.; E, Pa, V, VI B. scoparius L.: Thscap, Med.-Ir. Anal.; Pa B. steri/is L.: Thscap, Paleotemp.; E, Pa, V

3.3. Vegetation The prevailing uniform climatological, geomorphological and edaphological conditions in the area under study seem to be the determinative factors in the formation of the vegetation cover. The vegetation of all the islets of Enousses island group is characterized by the dominance of phryganic communities. These communities are characterized by the presence of FLORA (1994) 189

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perennial plant species, such as: Sarcopoterium spinosum, Cistus eretieus, Pistaeia lentiseus, Centaurea ~pinosa, Erica manipulijlora, Spartiumjunceum, Calieotome villosa etc. These perennial species combined with each other and other annual ones, contribute to the formation of a few vegetation types which differ mainly with regard the dominant species and the degree of plant coverage. Considering the factors which could have played a role in the observed vegetation diversity within the different islets or from islet to islet, the following ones could be referred: a. Random factors as discussed by RUNEMARK (1969) b. Human activities (agriculture, grazing, fires, etc.) c. Microecological differences. The differentiation of the microecological conditions which seems to be "poor" in the interior of the islets is quite obvious at the coasts where two distinct vegetational zones (littoral and sublittoral) can be distinguished as a result of the surf and the wind (RuNEMARK I.c.) Even though these zones are not very characteristic - regarding their floristic composition - on the studied islets, the vegetation types inhabiting the different zones are described below:

3.3.1. Littoral vegetation zone The littoral vegetation zone is characterized by a loose floristic composition accompanied by a low number of plant species and percentage of vegetation cover up to 30%. This zone is usually restricted to a narrow rocky belt since the geomorphology of the islets is not favourable to the formation of extensive sandy beaches. Sandy beaches of limited extent have been noted only on the islets Vatos and Enoussa. The main species participating in the plant cover of the sandy beaches are: Limonium virgatum, L. graeeum, Euphorbia peplis, Dittrichia viseosa, ]uncus heldreichianus, Leontodon tuberosus. The corresponding and dominant in our area, rocky plant communities are composed of the following species: Limonium graecul1l, L. gmelinii, Atriplex halimus, Centaurea spinosa, Crithmum maritimum, Helichrysum stoechas. Species like Narcissus serotinus, Sarcopoterium spinosum, Calicotome villosa have also been observed as participants to the floral composition of this vegetation unit in Vatos islet.

3.3.2. Sublittoral vegetation zone It constitutes a transitional zone situated between the littoral and the interior one occuring more or less in all the studied islets. Centaurea spinosa is the dominant plant species of this zone and associated with Pistacia lentiseus and Calicotome villosa, forms plant communities of a characteristic physiognomy. Often, this is a narrow zone located in a small distance from the sea but in some cases (Panaghia islet) its width is considerably expanded and then C. spinosa penetrates the interior zone forming an almost pure plant association. 76

FLORA (1994) 189

A special type of sublittoral vegetation occurs on the V atopoula islet, the smaller of the group, which due to its small size, is uniformly affected by the surf and the winds and it is not used by people for any activity. In this case the whole area of the islet (except the littoral rocky and erosed coasts) is characterized by a uniform vegetation, rich in species with a high degree of coverage (90-100%). The physiognomy of this plant cover is determined by the predominance of perennial shruby species like Centaurea spinosa, Pistacia lentiseus and Calieotome villosa. Various halophytic taxa such as Limonium graeeum, Limonium gmelinii, Silenefabaria, Maleolmia ehia, Lotus cretieus, Plantago spp. are also scattered all over the islet. Other plant species participating in the plant coverage of the islet are: Sareopoterium spinosum, Spartium junceum, Cistus spp., Phagnalon graeeum, Heliehrysum stoechas, Parapholis ineurva,Asparagus aeuti/olius, Dactylis glome rata etc.

3.3.3. Interior vegetation zone This zone appears on all the islets, except Vatopoula, occupies the greatest part of their area, and is inhabited by phryganic plant communities. We may distinguish the following three vegetation types based on the dominant species occuring on each one (physiognomical criteria). a. Type with Cistus creticus It is characterized by the intense and constant presence of Cistus cretieus which occasionally forms plant communities in combination with species like Sareopoterium spinosum, Pistaeia lentiscus, Calicotome villosa. In the interior of the islets Vatos, Panaghia, Pontikos, and in a restricted area of the western part of the main islet Enoussa, there exists a plant community composed of Cistus eretieus and Sarcopoterium spinosull1 as dominant species, and several other companion species (lower frequency of occurence and lower coverage degree) such as: Calicotome villosa, Pistacia lentiscus, Lavandula stoechas, Spartiumjunceum, Anthyllis herll1anniae, Satureja thYll1bra etc. In certain parts of Panaghia, Pontikos and Enoussa islets occurs another plant community, characterized by Cistus creticus and Pistacia lentiscus. These phryganic communities have a special aspect because of the occurence of well developed individuals of Pistacia lentiscus and Calicotome villosa, mainly in protected hollowed places. At the NW part of the Enoussa islet individuals of Pistacia lentiseus exceed the height of 2 m, and in neighbouring places, small stands of well developed individuals of Vitex agnuscastus have also been noticed. It is remarkable that in certain places - as on Pontikos islet - there appear "real mixtures" of plant communities predominated by Cistus eretieus, Sareopoteriull1 spinosum and Pistaeia lentiscus. The observed differentiation ofthe Cistus creticus - Sarcopoterium spinosum community on Vatos islet, due to the participation of Lavandula stoeehas, is also noteworthy. We have also to notice that Pistaeia lentiscus and Calicotome villosa participate in lower degree of occurence.

~-

b. Type with Sarcopoterium spinosum This vegetation type occurs mainly on the Enoussa islet on previously cultivated areas where agriculture is now abandoned. The physiognomy ofthese communities is given by the almost exclusive presence of Sarcopoterium spinosum. The percentage of plant-cover ranges from 60 to 80% and the mean vegetation height is ca. 30 cm. Pistacia lentiscus and Calicotome villosa are the species which participate with a low frequence in the floral composition of this community. According to RECHINGER (1951), two years of abandonment of previously cultivated lands is considered as sufficient for the development of a dense "carpet" ofSarcopoterium spinosum. A similar type ofphrygana has also been noticed in limited areas of Pontikos islet.

c. Type with Erica manipulij70ra This vegetation type occupies extensive areas in the western part of Enoussa islet and it is established on slopes with an inclination of 30 - 40%. It is characterized by high percentage of plant cover and mean plant height of 70 - 100 cm. We may distinguish two types of Erica formations: the one is characterized by Erica manipuiij7ora, Cistus creticus and the other by Erica manipulijlora, Lavandula stoechas. The floristic composition of both these Erica formations is completed by some more species such as: Cistus salviifolius, Calicotoll1e villosa, Pistacia lentiscus, Urginea maritima, Anthyllis hermanniae, Hypericum perforatum, Hyparrhenia hirta, Dactylis glome rata etc.

4. Discussion As has already been mentioned, the Aegean islets of the Enousses group were quite unexplored from a floristic point of view. Consequently, all the botanical information given in the present paper is considered as new records for the area. The Mediterranean character of the flora of the studied area is evident from the high proportion of the Mediterranean elements (East Mediterranean and Mediterranean s.l.) and of the Therophytes. In the flora of the area studied noteworthy is the absence of chorological elements with a distribution range restricted to the Eastern Aegean area and the neighbouring coasts of Asia minor. The absence of species presenting such a distribution pattern has already been revealed by studies in some other eastern Aegean islands (Chios: MEIKLE 1954, Psara: GREUTER 1976) and has been attributed to the recent geographical isolation ofthese islands. But ifthis is true for the islands of Psara and Chios with an altitude of 531 m and 1230 m respectively, then it is even more expected for our area. In the Enousses islets the highest hill does not exceed the altitude of 200 m and the shallow sea channels separating them from the adjacent coasts of Asia minor and Chios island minimizes even more the role of the isolation as a floral differentiation factor. Bearing in mind that the uniform geomorphological and climatological conditions prevailing in our area also do not favour the differentiation ofthe flora, then the observed floristic and phytogeographical banality should be expected. Moreover, the human activities lasting from ancient up to the present times must be considered as factors which have contributed to the disturbance of the natural ecosystems, the banality of the flora, and the more or less uniformity of the vegetation in our area. From historical reports it is also known that the main islet of the group (Enoussa = Oinousa) was inhabited since ancient times and vineyards covered the largest area of the islet. Today vineyards and other relative agricultural activities have been restricted to a few patches, but the opposite seems to be the case with the stock raising which still takes place in the main islet and in some smaller as well. Stock raising affects flora and vegetation by both overgrazing, and fire, since shepherds used to burn the area, in order to favour growing of pasturage species. We have already mentioned that

secondary vegetation types (e. g. Sarcopoteriull1 .lpinosul71type) are not uncommon in the studied area and there is also some evidence that not only the vegetation but also the floristic composition has probably been affected by human activities. This evidence deals with the representation of the Leguminosae and of the Therophytes, in each islet flora. In our area, especially from the data of Tables 1-3, it is evident that the Leguminous taxa and the Therophytes are represented by higher percentages in the floras of the bigger islets of the Enousses group (i.e. Enoussa and Panaghia). In these islets due to their size and geomorphology, a higher degree of human interference (agriculture, over-grazing, fires) is obvious. It is also necessary to point out that the increase of both Leguminosac and Therophytes in a local flora can be considered as relative index of disturbance for Mediterranean ecosystems. The postfire development of an impressive Leguminous flora has been noted several times and it is believed that underthese conditions Lelguminous plants act as a mechanism of replenishing esse"ntial nutrients - particularly nitrogen - which are dramatically consumed by fire (NAVEH 1974, ARIANOUTSOU & MARGARIS 1981, ARIANOUTSOU & THANOS 1992). Moreover, the high proportion of Therophytes, - which characterizes the flora of the bigger islets of the Enousses island group (Enoussaand Panaghia) - is also attributed to human activities according to Barbero et al. (1990). It has been noticed that the increase in grazing pressure throughout the southern mediterranean ecosystems leads to the occupation of the understories of the forest ecosystems (regardless of the altitude or type of ecosystem) by invasive Therophytes and indicate hyperdegradation (forest therophytization) (Barbero et al. I.c.). Taking into consideration the previous aspects it is remarkable that comparing the floras of the islets Enoussa and Panaghia, more affected by human interference seems to be the latter islet which is smaller and uninhabited. Such an observation should be explained only by taking into consideration that in such small and unhabited islets the different human activities (fires, grazing, hunting etc.) probably take place under quite uncontrolled conditions. The aspect that human activities lead to the disturbance of FLORA (1994) 189

77

the flora and vegetation of the ecosystems in the studied area is further supported by the rich and diverse flora of the Vatopoula islet, which is the smallest of our group. This islet, because of its small size (not more than 100 m in length), lying very close to the Vatos islet, does not favour agricultural or stock farming activities. The flora of this islet differs from those of the bigger ones (Enoussa and Panaghia) not only regarding the percentages of the Leguminosae and Therophytes (Table 3) but also with regard to some other floristic indices. It is noteworthy that in Vatopoula the East Mediterranean and the "Widespread" elements are equally well represented. Moreover, the flora of the islet is characterized by several species (Silene fabaria, Ephedra campylopoda, Malcolmia chia, Eryngium

mantunum, Plantago bellardii etc.), which have not been found in any other islet of the group. Since it is well known that floristic peculiarities, like those mentioned in Vatopoula, are common in most of the Aegean islets (RUNEMARK 1969), special attention should be given to the study and to the conservation, "in situ", of these ecosystems. Floristic elements of these ecosystems are very important for the knowledge of the total floristic diversity and of the species distribution pattern in the Aegean area. Moreover the small islets ecosystems could be used as basis for comparative ecological studies in order to understand the role of the human interference in ecologically similar habitats of the bigger Aegean islands.

5. Acknowledgements We gratefully acknowledge the assistance of the Greek Ministry nf Research and Technnlogy for financial support. The authors also thank

Dr. D. CHRISTODOUI.AKIS for his critical comments regarding the study of the vegetation.

6. References ARIANOl;Tsou-FARAGGITAKI, M., & MARGARIS, N.S. (1981): Producers and the tIre cycle in a phryganic ecosystem. In: MARGARIS, N.S .. & Moo:-JEY, H.A. (eds.): Components of productivity of Mediterranean-climate Regions. Basic and applied aspects. Dr. W. Junk Publishers, The Hague, 181-190. - & THANOS. K. (1992): Legumes in the post-fire plant communities of the Mediterranean ecosystems. International Workshop on the Role of Fire in Mediterranean Ecosystems. Bangyuls-sur-mer, France, Sep. 21-25. Book of Abstracts, p. 41. BAGNOLa.s, F., & GAUSSEN. H. (I <)57): Les c1imats biologiques et leur classification. Ann. Geogr. no 355, LXvie annee, 193-220. BARBERO, M .. BO!,;I)J, G., LOISEL, R., & Ql;EZEI., P. (1990): Changes and disturbances of forest ecosystems caused by human activities in the western part of the mediterranean basin. vegetatio 87: 151-173. CARI.STRilM, A. ( 1989): A survey of the tlora and phytogeography of Rodhos, Simi, Tilos and the Mannaris peninsula. Ph. D. TIIESIS. University of Lund. CHRISTODOCLAKIS, D. ( 1986): Flora and vegetation of the island of Samos. (Ph. D. Thesis. University ofPatras - Greece (in Greek with a German summary). DAVIS, P.H. (ed.) (1965-1985): Flora of Turkey & the East Aegean Islands. Vol. 1-9. Edinburgh Univ. Press. E~BERGER, L.c. (1955): Une classification biogeographique des c1imats. Rec. Trav. Lab. Bot. Geol. Zool. Fac. sc. de Montpellier.

78

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GRELTER, W. (1976): The Flora of Psara (E. Aegean Islands, Greece), an annotated catalogue. Candollea 31: 1<)1-242. - , BURDET, M.M., & LONG, G. ( 1984-1989): Med-Checklist. vols. 1,3,4. Geneve. MAVROMMATlS, G. (1980): Le bioclimat de Grece. "Dassiki Erevna" suppl. 1-63 + annexe. Athenes. MEIKLE. R.D. ( 1954): A survey of the flora of Chi os. Kew Bulletin 1: 851<)9.

NAVEII,Z. (1974): Effects of tIre in the Mediterranean Region. In: KOZI.OWSKI, T.T., & AlIl.CiREr\, C.E. (eds.): Fire and ecosystems. Academic ."Press, New York, 401-434. PIGMTTI. S. (ed.) (1982): Flora d \tali a, 1-3. Bologna. RAn,KIAER, C. (1934): The life-forms of plants and statistical plant geography. Oxford, Clarendon Press. RECIIII>CiER, K.H. (1943): Flora Aegaea. Florader Inseln und Halbinseln des Agaischen Meeres. Denkschr. Akad. Wiss. Wien, Math.-Naturwiss. KI.

105 (I). - ( 1951 ): Phytogeographia Aegaea. Denkschr. Akad. Wiss. Wien Math.-

Nturwiss. KI. 105 (2, 2). M. (I <)69): Reproductive drift, a neglected principle in reproductive biology. Bot. Notiser 122: 90-129.

Rl:!,;E~ARK,