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1nr.J. WildlandFire 6(2): 77.82.1996 BIAWF. Printed in U.SA.

Legumes in the Fire-Prone Mediterranean Regions: an Example From Greece*
Margarita Arianoutsoul and Costas A. Than&
'Department of Ecology and Systematics, School of Biology, Faculty of Sciences University of Athens, Athens 15784, Greece
Tel. 301 7284352; Fax. 301 7243325; E-mail

2Department of Botany, School of Biology, Faculty of Sciences, University of Athens, Athens 15784, Greece
Tel. 301 7284655; Fax 301 7234136; E-mail

Abstract. Mediterranean climate ecosystems of the Mediterranean rim are rich in legumes,both as woody shrubs of the late successional stages and as short-lived, pioneer herbaceous plants. In a survey of the floras of numerous geographic regions of Greece, legumes in Mediterranean partsare found tocontributeconsiderably more (1 1-16%of the total number of species) than in non-Mediterranean regions (6.8%). Legume presence is also notable in other Mediterranean-type ecosystemsof the world; nevertheless most leguminous taxaof the Mediterranean basin are herbs in contrast to asignificant contribution of shrubsin California and Chile. Due to their long-lived soil banks of hard coated seeds herbaceous legumes are usually among the early post-fie colonizers in the Mediterranean and may play a significant role in the succession of fire-pmne communities. Keywords: Mediterranean ecosystems; Greece; Leguminosae;
Ecological success

Ecological Success of Legumes

Introduction The legume family is considered an ecologically successful family of flowering plants worldwide (Rnndel 1989). Legumes occur almost in every terreshial hiome, ranging from 80 m tall giants in the forests of South East Asia, to small desert ephemerals with a life span of a few months. The legume family comprises more than 17000 species which correspond toapproximately 7%of thecurrently described vascular plants of the world. It is the third family in absolute number of taxa, exceeded only by Compositae and Orchidaceae (Mabberley 1987). In the Mediterranean rim and in Greece it is the f i s t in the number of taxa it has (Tutin et al1964-1980). while it is high in the rank in other Mediterranean climate areas of the world, as for example, in the Cape Floristic Region, where it is the fourth (Cowling and Holmes 1992). *Originally presented at the 2nd International Conference on Forest FireResearch. November 21-24.1994, Coimbra, Portugal.

The legume taxa are ecologically very successful in terms of dominance and productivity. The morphological and physiological adaptations utilized by legumes are not unique. Other plant families demonstrate similar adaptive flexibility for one or several traits, but none exhibits the tremendous range of adaptations shown by legumes. These traits are expressed through remarkable levels of adaptive variation in morphological characters such as growth form, canopy architecture, mot architecture, leaf morphology, pod and seed sbuchlre and in physiological features such as phenological controls, solar tracking, water relations, hard seed coats, nitrogen fixation and mycorrhizal associations (Rundel1989). In more detail, leguminous plants turn the leaves so that to have their surface perpendicular to the sun rays in the morning and the afternoon in order to maximize the incident radiation, while they keep them parallel to the sun rays at midday, in order to reduce the excessive heat load. They employ stem photosynthesis, which in certain cases may amount to about one third of the leaf tissue photosynthesis, with high water use efficiency of the fixed carbon (Rundel1989). A similar high water use efficiency is presented by the phyllodes which are often developed as a sclerophyllous substitute of leaf tissue (Rundel 1989). An efficient water economy is also enhanced by adaptations such as nyctinastic closureof leaves and the phreatophytic habit in many desert legumes e.g. as the Prosopis glanduloso (Nilsen et al1981). As far as it concerns nutrition, the adaptive mechanisms they have are related to nitrogen Fixation through the development of nitrogen fixing root nodules in soils with limited nitrogen availability and the formation of mycorrhizal associations which enhancephosphorus uptake. Finally, they adopt various dispersal modes (endozoochory, autochory etc); in addition, the almost ubiquitous presence of hard seed coats ensures maximum seed survival (when fruits are consumed by animals) and contributes to the formation of permanent, long-lived soil seed banks.

1964-1980). while with increasing dmught interactionsthe dominant growth forms become reduced in stature and leaves become reduced in size and have lower longevity.3% for the sclerophyll forests of South Australia (Specht 1972). Davis et al. which covera bmad range of precipitation regime and length of dry periods. 1984-1989)are found in Greece. One hundred per cent (100%) of the legume flora in the tropics of SE Asia are - Table 1 . The cold temperate environments typically have a relatively low diversity of legumes. Hopper and Maslin 1978. The existence of tall mountains is also considered to be the reason for the rather low percentage values of several areas with overall Mediterranean climates (5-13. stiped bars). simply because they are located around the Mediterranean Sea in addition. while in the Neotropics the frequency of legumes in the see flora is much higher. Malaisse 1978). 1984-1989. with typical values rangingbetween 11-13%. 1988.4% legume percentage (Hulten 1968).Boucher1977. southern parts of North African countries. (Table 2). When gradienu in legume growth form and phenology are viewed from the wet Wpical forest to dry deserts and Mediterranean climate ecosystems. Lathrop and Thome 1978. 16% of the total flora are legumes.g. In the cold and dry White Mountains of California only 4. 416 legume species are recorded for Greece (Greuter et al. which do not have aMeditemeantype climate. Mountainous areas (dotted bars 1-4) have a significantly lower contribution (6-7.8-13. almost 49% of the species belonging to theLeguminosaefamily and referred in Flora Europaea are found in Greece (Tutin et al. although they are considered as such. Estimations based on the legumes listed in the MedChecklist (Greuter et al. Hoover 1970.7-9: Levyns 1966. in the Med Checklist.8% ofthe Chilean mammal (Rundel 1981) to relatively high. Bond and Goldblatt 1984. The flora of Alaska includes a similar 4. 13.3% of the flora are legumes (Loyd and Mitchell 1973). C. whiie in the deserts of Central Australia legumes comprise 13% of the flora (Jessop 1981). CHILE (6) . tall mountain ranges in all countries).b:Rundel1981. there are clear patterns of change (Rundel 1989).596) belong to the mountain flora (Strid 1989). most notably with species of Prosopis and Acacia (Gentry 1942).3 . The Sonoran Desert has 11. while only herbaceous forms of legumes can be found in Alaskan flora. Contribution of leguminous mxa to the florasof scvcrd Mediterranean climate regions of the world. Mooney 1977. Regions 5 and 11 are not mountainous hut legume contribution is presumably decreased by the halophytic vegetation present (which is generally underrepresented in legumes). (1-5: Thomas 1961. SOUTH AFRICA (7-9) AUSTRALIA ( 1 0 ) ' ' 8. with evergreen and semi-evergreen leaf phenology. whiie in the subtropical forests of Brazil and Africa their participation ranges between 15 and 19% in ascending order (Eiten 1972. In more temperate environments. Growth forms change fist to lower stature eees and shrubs and fmally to dominationby low shrubs and herbs. However. this is an underestimation. Greuter 1991) give for the legume flora of the entire Mediterranean Basin an overall value of 7. The floristic diversity of legumes in the Sahara Desert is similarly high (Ozen& 1958). Distribution of Legumes Legumes are abundant and frequently dominant in diverse terrestrial hinmes. region 5. Mediterranean climate ecosystems of the world have legume diversities ranging from very low values of 3. 1984-1989). Legumes in the Mediterranean plant communities According to estimations the total flora of Greece is composed of 4700 species. 10: Specht 1972). Ullmann 1985). typically M e d i m e a n regions of Greece (Figure 1. Approximately 23% of the 1833 species of Leguminosae listed in the Med-Checklist (Greuter et al. herbaceous perennials become the dominant growth form (Rundel 1989). In the dry forests of the west coast of Mexico. Region (Reference) Legumes %of h e 1-1 flm . Whibley 1980. Considering that the mountainous flora represents 42.A. since countries. In the tropical forests of Africa legumes comprise 10% of the total flora (cited by Rundel 1989). e s e r t of California has Deep Canyon in the Sonoran D a legume flora of 11-13% of the total flora (Zabriskie 1979). in altitudes higher than 1800 meters. northern and Atlantic France. (Table 1). speciesof Acacia are overwhelmingly dominant in these communities (Palgrave 1977. In a survey of the floras of numerous (30 in total). One hundred and eighteen of these species (2A. larger or smaller parts of the circum-Mediterraneancountries are not truly Meditemnean (e. much higher than the overall valuesof Greece (orcypms).1% of the total flora of Greece. trees. grey bars numbered 14-43) it is shown that legume percentage conaibution in the total flora ranges from 11 to nearly 16% and in the majority of the cases between 12-14%. and Thanos. it is evident that the legume flora is considerably underrepresented in the mountain flora of the country. while in North and South America woody legumes may make up to 90% or moreof the woody plantcover in thornscrub. At more mesic sites (seasonal tropical forest) legumes are predominately trees and woody lianas.5%). (like Libya and Bulgaria) are included in the list.6%legume flora. In Africa and Aushalia. Similarly.6%. In the dipterocarp forests of South East Asia legumes comprise only 3-5% of the total tree flora..

8 6. (Tzanoudakis 1981) 23 Skiathos Isl. Euboeic Gulf islands (Sarlis 1981) 37 Elaphonesos Isl. (Cantargy 1889) 9 Euboea Isl. (Phitos 1960) 14YiouraIsl. (Tsimburla and Yannitsaros 1992) 39 SkantzouraIsl.3 Countries Greece Mmtains of Greece Crele 4700 1980* 1586 1800 7. 1990) 30 Chios Isl. (Economidou 1973) 19 HymetmMt. (Phitos and Damboldt 1985) 13 Central Euboea Isl. seasonally covered in part by sea waves. 1991) 27 Aetoloacarnanian lakes (Koumpli-Sovantzi 1983) 28 Cassandra Peninsula (Lavrentiadis 1961) 29 Dionysades Isls (Christodoulakiset al. The high legume contribution though is attributed to the human intervention and particularly the overgrazing pressure which is postulated to favour legumes.0 9. (Christodoulakis 1986) 34 Kos-Kalymnos-Pserimos-Telendos Isls (Hansen 1980) 35 Spetses Isl. and 15 islets (Raus 1989) 16 CorfuIsl.9 9. ~trofiliacoastal wetland. (Rechinger 1961) 10 Vertiscos Mt. (Kamari et al. 18. range (Pavlidis 1982) 11 Strofdia hydrobiotope (Gwrgiadis et al. is mainly a marshland. * (Meikle 1954) 31 Santorini Isls (Hansen 1971) 32 Sithonia Peninsula (Pavlidis 1976) 33 Samos Isl. (Yannitsaros 1971) 38 Piperi Isl.6 8. (Economidou 1969) 24CytheraIsl. Floristic region Total number of species - Number o f legume species Legume cmuilutian ( 9 6 ) EUW Mediterranean 11557 2 4 0 W 844 1833 416 118' 7. Evstratios Isl. (Greuter 1976) 46 Oinousses Isls (Panitsa et al. (Georgiadis et al. 4 ~ r e s p a s ~ a kNat.Contribution of legumes to the total flora of various geographic regions of Greece (m the cases marked with * the contribution dus subs~ecies basis). (Greuter 1979) 45 PsaraIsl. 198b) 6 Cholomon Mt. while region 11.3% (Panitsa e t al. (Voliotis 1967) 7 CreteId. in particular). As far as it concerns regions 44 and 45. (Zerlend'is 1973) 20 Ag. (Papatsou 1975) 41 Paxi Isl. 1990) 12 Cephallonia Isl. mikolaidou and Yannitsaros 1992) 36 S.Contributionoflegumes m the floras ofselected floristic 79 regions.3 Cyplus 157 168 * Numben refer to species plus subspecies Psathoura. (Yannitsaros 1969) 25 Mt. Region 25 is a mountainous area with a climate between those of Meditemnean and Central Europe. Dodecanesus (Panitsa and Tzanoudakis 1991) 44 Kastellorizo Isl. which was performed during spring-time. (Sarlis 1994) 18 SkopelosIsl.Legumes in the Fire-Prone Mediterranean regions Table2. Paikon grazelands (Drossos and Athanassiadis 1989) 26 Kira Panagia Isl. Figure 1. is a small rocky island. (Gustaffsonand Snogerup 1974) 40 Nisyros Isl. the extreme values may be the artifact of the plant list compilation. (Barclay 1986) 8 Lesbos Isl. This same argument i s used in the case of Oinousses islands (region 46) to explain the dramatically high percentage. 1988) 15 ArmathiaIsl. 1994). e s Park (Pavlidis 1985) 5 ~sakuralsl. (Georgiou 1988) 17 Syos Isl. (Snogerup 1991) 21 Schinias pine forest (Bmfas and Karetsos 1991) 22 Yiaros Isl. 1 Mountains of Greece * (Strid 1991) 2 Lailias Mt. thus favouring legumes and not including several autumn and winter flowering plants (geophytes. Noliotis 1977) 3 Macedonian has been estimated ona s~ecies Mts (Quezel and~ontanhriopoilos 1968.. . (Snogemp et al. 1986) 42 Kassandrapine forests* (TS~~SON and Karagiannakidou 1987) 43 Seven islets of N. (~nog'erupet al. 1994).

) 103(1O~pp. Armstrong. Thanos et al1996). and P.) 22. while 75. Doussi and Thanos 1994). Checklist ofthevascularplantr. Fire-followers in San Diego County. which is seed germination. Papavassiliou et al 1994.6 (638 spp. Table 3. Bond. Mooney). UK. (Naveh 1967. shortage of available water). only 3 (0.A. Cowling).. ENVIRONMENT) for financial support. References Arianoutsou-FaraggitakiM.. the herbaceous elements dominate. Although detailed dataabout the floristics and the dynamics of this succession are scarse there are cerrain evidences that an enrichment of the local floras in leguminous elements occurs. 1992.6 (5 spp.) 22.Aprovisionalchecklistof floweringplans and ferns in the Cape Hangklip area. RM. The Hague.S. but also requires the high temperahlKS developed by fire for the induction of germination (Doussi and Thanos 1993. which does not only permit the survival of seeds during a wildfire.C. It is weU known that seeds of legumes possess a hard. but. Greece. where legume taxa constituting approximately 9% of the flora of a mature stand more than doubled immediately after fire (Kazanis and Arianoutsou 1994). their prolific presence in theregenerating postf i e communities has been attributed to their ability to fix atmospheric nitrogen. Mareark and I I A. Most of these plants are annual or biennial herbs (op. water impermeable seed coat. Flora andvegetation. Most herbaceous legumes overcome fire occurrence through seed germination. approximately 90% of the relevant flora of Pinus halepensis forests of Attica. project 91 ED 944) and to the European Union (PROMETHEUS projectno EV5V-CT94-0482.5%) of U 13 species the Papilionoidae (Lotoideae) subfamily. o~s~N. C. 1977. pages 181-190. thus competing with other non-leguminous plants in the early post-fire environment.1 (I8 spp. to really impressive. and Thanos.80. The authors express their gratitude to theGeneralSecretariatofResearchandTechnology.M. Producers and the fne cycle in a phryganic ecosystem. P. 1964-1980).6 (188 rpp. There is strong evidence that the majority of the legume plants of the floras of several Mediterranean regions of Greece are annual.1% are @ees. in the leguminous flora of Europe only 2.5% of the legume flora of Samos island (Pinus brutia forest and maquis). Boston. biennial or perennial herbs: 85. The prolific appearance of the leguminous plants in the burned areas lasts for only the very early succesional post-fm stages (Kazanis and Arianoutsou 1994). which gradually become restricted both in species represented as well as in cover.c). Arianoutsou-Faraggitaki and Margaris 1981). A belonging to Caesalpinioideae and Mimosoideae are trees. One of the most potentially adaptive traits of these plants is increased post-fie seed germination. In: The Ecoloev of Fvnbos.) 2. while woody ones may also be resprouters (pers.) 76.1973). Growth forms in the European legumes. mature stage). A descriptive catalogue. Although f i e is considered as aclimate related hazard. Plants of the Cape flora. The characteristics of this recruitment are strongly selected for at "normal" fire intervals. C. Legumes have been referred as an important component of post-fire successional communities of Mediterranean climate regions (Naveh 1967. Goldblan. The vegetation of the Mediterranean communities is extremely flammable during summer months. 1989. Holmes. because of the accumulation of great amounts of dead material (standing and fallen as litter) and the prevailing climatic conditions (high temperatures.S. Fire and Diversitv . London. the actual effectiveness of this nodulation still remains to be studied. as in the case of a Californian chaparral (Armstrong 1977). 61 %Trees % Shmbs %H e r b s 1 W (3 spp. W. pages 23- According to Flora Europaea (Tutin et al. Kazanis and Arianoutsou 1994. all growth forms are represented. Oxford University Press.3 (188 spp. 1981. Overall. D r W..Nutrienu.) 75.). . On the contrary. 1986. (Kazanis and Arianoutsou 1994). recently burned. It is not surprising therefore that the plant species of the fire-prone environmentsof the Mediterranean ecosystems display many adaptations to f i e (Naveh 1973.6% are herbaceous plants Vable 3). This habit might be related to their most common adaptation towards fire.8 (638 spp. (Christodoulakis 1986). Up to the present.Englera 6:l-138. and P. 76.Greece (PENED. (edircd bv V. Journal of South African Botany 4357. M. In: Corncanens uf Producuviy of ~edit&anem T & ~cosystem\:Basic and Amlied .P.andN. as in the case of a Pinus holepensis forest in Attica. 1984. Junk Publishers. however.M.4%) belong to Caesalpinioideae subfamily and 10 (1. Subfamily Caesalpinioideae Mimoroideae Lotoideae - . it is also an environmental element itself. Thanos et al. out of 844 legume species native in Europe. Acknowledgments. Papavassiliou and Arianoutsou (1993) have found that post-fire legumes do appear to have nodhation capacities. Rundel 1981.9. (edited ~ by R.80 Arianoutsou. in the Papilionoideae subfamily. the work published cannot answer the questions why legumes are so abundant after fire and why legumes are so dominant in the burned places during only the early post-fle successional stages.) 0. Joumal of South African Botany 13Supplemenmy Volume:1455 Boucher. Papavassiliou and Arianoutsou 1993. Asmu. Arianoutsou-FaraggitaJci and Margaris 1981.Crete.Margaris. Cowling. This enrichment may be slight. On the other hand.9% of the legume flora of Kythira island (mostly phryganic vegetation.) LEGUMINOSAE Barc1ay. Frernontia 43. (Yannitsaros 1969). 1977.8%) to Mimosoideae in contrast to the 831 species (98.

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